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DEEL 117 MUS. COMP. ZOOL 1974
i LIBRARY

MAK 1 2 1975

HARVARD
UNIVERSITY

TIJDSCHRIFT
VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

| Tijdschrift voor Entomologie, deel 117, 1974 |

Afleveringen 1-2 verschenen 4.VI.1974
Afleveringen 3-4 verschenen 23.VIII.1974
Afleveringen 5-6 verschenen 15.X.1974
Aflevering 7 verscheen, 31.X11.1974

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INHOUD VAN DEEL 117

CHVALA, M. J. Doskoür, J. H. Mook & V. Pokorny. — The genus Lipara
Meigen (Diptera, Bu systematics, morphology, behaviour, and
CODE ss nt. . AAN A IO en A 1

Doskotit, J., zie CHVALA.

Hure Ris LAMBERS, D. — On American Aphids, with descriptions of a new genus
and some new species (Homoptera, Aphididae). . . . . . . . . . 103

Jong, R. DE. — Systematics and evolution of the Palaearctic Spzalia species (Lepi-
hot, HER Ne EE 0 0. 225

LiEFTINCK, M. A. — Review of Central and East Asiatic Habropoda F. Smith
with Habrophorula, a new genus from China (Hymenoptera, Anthophoridae) 157

LitH, J. P. VAN. — Revision of the Psenini of the Ethiopian Region, including
Malagasy (Hymenoptera, Sphecidae, Pemphredoninae) . . . . . . . . 39

Mook, J. H., zie Chvála.

MUNROE, E. — Pyralidae of the Third Archbold ee IE Ma Odon-
Bimaenkenidoptera) 2 e... li : PA 27

Pokorny, V., zie Chvala.
RossEM, G. VAN. — The Gravenhorst, Schigdte and Foerster types belonging to
the genus Megastylus Schigdte, 1838, with keys to the Be a

Ichneumonidae, Microleptinae) . . . . . . : LT

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TIJDSCHRIFT Sr
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UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

x M. CHVALA, J. Doskoär, J. H. Mook & V Pokorny. — The genus Lipara Meigen
a | (Diptera, Chloropidae), systematics, ae behaviour, and ecology, pei 1-25,
EE 1—22, Pl. 1—7.

| Tijdschrift voor Entomologie, deel 117, afl. 1

. Gepubliceerd 4-VI-1974

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THE GENUS LIPARA MEIGEN (DIPTERA, CHLOROPIDAE),
SYSTEMATICS, MORPHOLOGY, BEHAVIOUR, AND ECOLOGY

M. CHVÁLA *), J. DOSKOCIL'), J. H. MOOK?) & V. POKORNY ©)

ABSTRACT

Four species of the genus Lipara, viz.: L. lucens, similis, rufitarsis and pullitarsis, occur in
Europe, where they cause galls in Common Reed (Phragmites communis). The specific differences
of eggs, larval, and adult stages are described, and a general account is given of the life cycle,
gall formation, precopulatory and ovipositional behaviour, habitat relations, and influence of
parasites and predators with emphasis on the known differences between the species.

CONTENTS

1. Introduction Md. 2
2. Systematics and morphology . ballate ie ein ad Alet hoy. sie 2
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ee VE re SER PO AR ER EEE De 24

1) Department of Systematic Zoology, Faculty of Natural Sciences, Charles University, Viniëná 7,
128 44 Praha 2, Czechoslovakia.

2) Institute of Ecological Research, Kemperbergerweg 11, Arnhem, the Netherlands.

3) Institute of Scientific and Technical Information, Czechoslovak Akademy of Agriculture,
Slezská 7, 120 56 Praha 2, Czechoslovakia.

2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

1. INTRODUCTION

The galls on Common Reed (Phragmites communis Trin.), caused by flies of the
genus Lipara, have attracted the attention of many generations of entomologists in
Europe. During more than one hundred years, after several entomologists in Vienna
had collected reed galls and studied their inhabitants (Schiner 1854, Heeger 1856,
Loew 1858, Giraud 1863), three species were recognized: L.lucens, rufitarsis, and
similis.

In 1957 Ruppolt mentioned the occurrence of two types of larvae of L. rufitarsis,
one in typical galls and the other in galls indentical to those of L. /acens. He supposed
that these might be two different species.

From 1967 to 1970, Pokorny took up the problem again in Czechoslavakia, and
found that the two types of L. rzfitarsis make distinctive galls. The differences he
observed also in the life cycle and the behaviour of the larvae were such that he expected
the insects to belong to two different species. He submitted the adults, bred from the
galls, to Doskocil and Chvála who indeed recognized them as two different species,
one being L. rufitarsis and the other, a previously undescribed species which they named
L. pullitarsis (Doskotil and Chväla, 1971).

In recent years Mook studied the ecology of Lipara lucens in the Netherlands (Mook,
1961, 1967) and also collected data on the ecology of L. rufitarsis and pullitarsis (L.
similis does not occur in the Netherlands). In co-operation with Miss C. G. Bruggemann
he studied the behaviour of the adult of L. /zcens (Mook and Bruggemann, 1968, 1971)
and also collected data on the behaviour of the other species. These data corroborate
the view that four different species are involved.

It seemed worth while to collect both the published and the unpublished data in order
to give a survey of the similarities and differences in the biology of the four species.
The results are presented here. The authors have each treated those aspects of the
biology with which they are most familiar, and they remain responsible for the relevant
section: Chväla and Doskoëil for section 2, Pokorny, for section 3 and Mook, for
sections 4 and 5. It should be kept in mind that the different geographical localities
where the authors collected their data may have influenced their findings.

2. SYSTEMATICS AND MORPHOLOGY
a. Adults
Genus Lipara Meigen, 1830

Lipara Meigen, 1830: 1. Type species, Lipara lucens Meigen, 1830 (monobasic).

Homalura Wiedemann, 1830: 573 (nec Meigen, 1826), junior homonym. Type species, ee
grisea Wiedemann, 1830 (monobasic).

? Gymnopoda Macquart, 1835: 503, subjective synonym. Type species, Gymnopoda tomentosa
Macquart, 1835 (monobasic).

The genus Lipara Meigen belongs to the family Chloropidae, subfamily Oscinellinae,
and comprises medium-sized to large blackish species, 3.3—7.3 mm, with dense pale
pubescence on mesonotum. Arista thin, not pubescent, antennae broadly separated at
base by a wide pollinose facial keel, very prominent in lateral aspect. Eyes rather small
and finely pubescent, face very deep, almost as long as one half of the eye-height;
vibrissal angles absent.

M. CHVALA, J. DoskoCr, J. H. Mook & V. Pokorny: The genus Lipara 3

The genus nearest related to Lipara is Calamoncosis Enderlein wich comprises dis-
tinctly smaller species, 2-3 mm, with equally oval head in profile, with facial “keel” only
indistinctly developed. In Lipara the facial keel is prominent and very broad over the
whole length, as far as the mouth cavity, or, if it is narrowed in the middle between
the antennae, again widening out below; viewed from the side, the keel is strongly
prominent above the antennae, as is the lower part of the frons, so that the head in
profile ist strongly prominent anteriorly.

The larvae of all Lipara species cause more or less distinct galls on stems of the
Common Reed (Phragmites communis Trin.).

The genus has been recently revised by Doskoëil and Chvála (1971) and as a result
four Palaearctic species (viz., lucens, similis, rufitarsis, and pullitarsis) were introduced
as essentially European species. Two of them, /zcens and similis, have been found
separately as immigrants in North America.

The four Palaearctic Lipara species may be keyed as follows:

1 Thoracic pubescence long, arranged in ee stripes, A a ridged

appearance. Large or smaller species . . RE 2

— Thoracic pubescence shorter, uniformly spread not angel i in Stripes Smaller
Speciesy I . . to sac 3

2 Thoracic eee: bes Helen uo species 5. Je 3 mm, gel keel very
progam Pa. .\. ee ee ens

— Thoracic pubescence ele Smaller Ser 3. 34. 6 mm, ia keel narrower
A similis

3 Facial keel a a Ato i Sii ol as banal as one-half of antennal
segment 3. Male genitalia with fused cerci. Length 3.3—5.0 mm pullitarsis

— Facial keel narrowed in the middle, widening above and below, less broad than
one-third of antennal segment 3. Male genitalia with paired cerci. Length 3.5—
ENE ee NN af ee 4 AN raftarsis

Lipara lucens Meigen, 1830

Lipara lucens Meigen, 1830, Syst. Beschr. 6: 1.
Homalura grisea Wiedemann, 1830, Aussereurop. zweifl. Ins., 2: 573.
? Gymnopoda tomentosa Macquart, 1835, Hist. nat. Ins. Dipt., 2: 502.

Diagnosis. A conspicuously large and robust species, 5.3—7.3 mm, the largest species
of the genus and family in the Palaearctic region. Thorax with long, brassy-yellow
pubescence on mesonotum which is arranged in distinct longitudinal stripes, producing
a ridged appearance. Antennae yellowish on basal segments, segment 3 mostly blackish.
Facial keel conspicuously broad and with almost parallel sides. Legs extensively darkened.
Male genitalia (Fig. 2) large, with small but rather widely separated, apically pointed
cerci.

Distribution. A widely distributed and everywhere common species in Europe, ranging
in the North from England through the Netherlands, Denmark, and southern Sweden,
eastward as far as the northern regions of the European part of the USSR, a very
common species in central and southern Europe, immigrant ín North America (Con-
necticut).

4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

Lipara similis Schiner, 1854

Lipara similis Schiner, 1854, Verh. zool.-bot. Ges. Wien, 4: 172.

Diagnosis. Closely related to /zcens, having the same type of long pubescence on
thoracic disc which is arranged in distinct longitudinal stripes, producing a ridged
appearance. In contrast to /zcens, the thoracic pubescence is distincly whitish to silvery-
grey; similis is a much smaller species, 3.3—4.6 mm. Facial keel rather broad, slightly
widened above and below. Antennae and palpi yellowish. Legs mostly yellow on tibiae
and tarsi, but in some specimens tarsi extensively darkened. Male gentitalia (Fig. 4)
small, cerci not very wide apart, small, and apically distinctly rounded.

Distribution. The species is rather common and widely distributed in Central Europe
(Poland, Germany, Czechoslovakia, Austria, and Hungary), and has also been recorded
from England, France, and the western regions of the European part of the USSR.

Figs. 1—8. Head in profile and male genitalia (caudal view) of Lipara: 1, 2, Iucens Meig.; 3, 4,
similis Schin.; 5, 6, L. pullitarsis Dosk. et Chv.; 7, 8, L. rufitarsis Loew. Scale = 0.3 mm

M. CHVALA, J. DoskoCr, J. H. Mook & V. POKORNY : The genus Lipara 5

According to Sabrosky (1958), the species was intercepted in the USA (Hoboken) in
Phragmites packing from a ship from the Netherlands. It was also mentioned for the
Netherlands by Docters van Leeuwen (1957), but his description and illustration of the
gall do not fit. The species has not been reared from Dutch galls (de Meijere, 1949).
Dr. Th. van Leeuwen, Dr. W. J. Kabos, Dr. G. Kruseman (Amsterdam) and Dr. S. J.
van Ooststroom (Leiden), who were so kind as to inform us of their experiences, also
did not find the species in the Netherlands.

Lipara pullitarsis Doskoëil et Chvála, 1971

Lipara pullitarsis Doskotil et Chvála, 1971, Acta ent. bohemoslov., 68: 102.

Diagnosis. A smaller species, 3.3—5.0 mm, with mesonotum densely covered with
mostly brassy-yellow short pubescence, not arranged in longitudinal ridges. Facial keel
very broad, the narrowest part in the middle about as broad as half the width of antennal
segment 3. Antennae mostly black, only basal segments sometimes very dark brown; palpi
black. Legs mostly black, extreme tips of femora and extreme bases of tibiae slightly
brownish; fore tarsi black, posterior four tarsi yellowish-brown on basal segments, apical
segments extensively darkened. Male genitalia (Fig. 6) rather large, with broad parameres
and entirely fused long cerci. The Central European and southern Scandinavian pop-
ulations have extremely darkened antennae and legs, but we have seen paler specimens
from the Netherlands that superficially resemble r#fitarsis, differing from the latter by
a much broader facial keel and quite distinct male genitalia.

Distribution. This species has long been confused with rwfitarsis, and it is highly
probable that many records on the distribution of r#fitarsis refer, in fact, to pullitarsis.
The species was described very recently from Czechoslovakia but we have also seen
documentary material from the Netherlands, Denmark, and Sweden.

Lipara rufitarsis Loew, 1858

Lipara rufitarsis Loew, 1858, Wien. Ent. Monatschr., 2: 57.

Diagnosis. A smaller species, 3.5-—5.3 mm, resembling pullitarsis in having the same
type of pubescence on thoracic disc; the hairs are short and not arranged in longitudinal
ridges but, in contrast to pullitarsis, the pubescence is distinctly silvery-grey and the hairs
on the hind part of mesonotum and on the scutellum are distinctly longer. Facial keel
is much narrower than in pullitarsis, widening above and below, the narrowest part in
the middle being hardly as broad as one-third the width of antennal segment 3. Antennae
reddish-brown except for outer side of segment 3 and arista which are extensively
darkened or almost black; palpi black. Femora and tibiae predominantly black, the
apical tips of femora, both rather broader tips of tibiae and whole tarsi yellowish- to
reddish-brown. Wings with cubital and discal veins almost straight, not curved as in
pullitarsis. Male genitalia (Fig. 8) rather large, with narrower parameres, long cerci
apically distinctly separated.

Distribution. L. rufitarsis is recorded in the literature as a common species throughout
Europe, but at least some of these records must concern pullitarsis. We have seen the
documentary material from the Netherlands, Czechoslovakia, Austria, and Italy, and

6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

according to Dr. Hackman (letter of 24.8.1971), the material from southwestern Finland
also belongs to rfitarsis. On the other hand, having revised in Copenhagen and Lund
all of the accessible vufitarsis material from Denmark and southern Sweden, one of the
authors (Chvála) comes to the conclusion that all of it belongs to pullitarsis.

b. Immature stages

There are few older data on immature stages of Lipara; a summary of all the records
up to 1950 is given by Hennig (1952). He found only four records of larva and
puparium of /zcens in the literature, but the record of larva by Vimmer (1925), must,
in fact, refer to rufitarsis judging from the Czech description and figures. Ruppolt
(1957) described eggs and larval stages of /cens. The most complete treatment of the
morphology of immature stages of lwcens is by Waitzbauer (1969). No other records are
available.

Most of the eggs, larvae, and puparia used for the study discussed in this section,
belong to material collected and bred by J. H. Mook and V. Pokorny.

Eggs

The eggs of the four Palaearctic Lipara species are very distinctive. In general, they
may easily be separated into two different groups, viz., (1) the /ucens type and (2)
the rufitarsis type. The former type (/ucens and similis) is characteristically rather
broader, more oval, and light yellow in colour. The differences between the eggs of
lucens (Plate 2b) and similis (Plate 2d) are very slight; the egg of similis is almost
as long and as wide as that of /cens, but the latter has more distinct sculpture, con-
sisting of rather deep longitudinal furrows (Plate 2a). The measurements of the eggs
of these two species are as follows: /ucens: (N = 40) length 1.37—1.69 mm (M =
1.57 mm), width 0.25—0.36 mm (M = 0.32 mm); similis: (N = 20) length 1.35—
1.46 mm (M = 1.42 mm), width 0.23—0.32 mm (M = 0.27 mm). The ratio of
length to width is about 5:1 in both /zcens and similis.

The second type of egg (rufitarsis and pullitarsis) is characteristically long and narrow,
and distinctly blackish in colour. The egg of pullitarsis (Plate 3b) is distinctly narrower,
slightly shorter, rather circular in cross-section, and the longitudinal furrows (Plate 3a)
are quite distinct. The eggs of rufitarsis (Plate 3d) is less slender but slightly longer,
somewhat triangular in cross-section (rounded above, flattened below), and the long-
itudinal furrows are less numerous and barely visible; the whole egg is therefore rather
smooth. The measurements of these two eggs are: pullitarsis: (N = 30) length 1.13—
1.26 mm (M = 1.21 mm), width 0.07—0.11 mm (M = 0.09 mm); rwfitarsis: (N
= 40) length 1.39—1.53 mm (M = 1.50 mm), width 0.15—0.20 mm (M = 0.18
mm). The ratio of length to width in rwfitarsis is 8 : 1, in pullitarsis 13 : 1.

Larvae
Larvae are elongate, light yellowish, with one or both tips more or less sclerotized;
the length of mature larvae varies from 5.5 to 12 mm; all measurements were taken
from larvae collected in the winter or early spring. The whole body is rather smooth,
integument exclusively with regularly spaced flower-like not sclerotized papillae and
sometimes with striae, especially on anterior segments. The sclerotization of anterior
and posterior segments, including the number of buds on anterior spiracle, seem to be

M. CHVALA, J. Doskocit, J. H. Mook & V. POKORNY : The genus Lipara 74

the best specific characters. Posterior spiracle consists of three simple radiating slits,
interstigmatal processes are branched (? absent in rzfitarsis).

The mature larva of /ucens (Plate 6a) is conspicuously large and stout, 8—12 mm
in length and 2—3 mm in width. Only anterior segments dorsally sclerotized (Fig. 9),
more heavily on segment 1 as far as the anterior spiracle, a slight sclerotization con-
tinues over anterior spiracle on dorsum of segment 2 and anterior half of segment 3.
Segment 1 sometimes also slightly sclerotized ventrally. Posterior margin of anterior
three segments on dorsum with small sclerotized papillae in a row; on venter also
present but hardly visible (not sclerotized). Anterior spiracle (Fig. 13) with 11 to 12
(exceptionally up to 14) buds, their number not constant on each side; the usual com-
binations are 11—11, 11—12, or 12—12, in rare cases, 12—13 or 12—14.

Figs. 9—12. Four anterior segments of mature larva of Lipara (lateral view). 9, L. lucens Meig.;
10, L. similis Schin.; 11, L. pullitarsis Dosk. et Chv., 12, L. rufitarsis Loew. Scale = 1 mm

8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

The mature larva of similis (Plate 6b) generally is smaller and especially more slender,
5.5—10 mm in length and 1—2 mm in width; it is the only species of the genus with
distinct sclerotization of both tips. Segment 1 (Fig. 10) heavily sclerotized dorsally,
except for a narrow anterior strip, segments 2 and 3 heavily sclerotized dorsally, almost
blackish, as is segment 1. Segment 4 dorsally with a slight sclerotization, venter of four
anterior segments pale, membraneous. Segments 5 to 10 membraneous, segment 11
slightly sclerotized dorsally and ventrally, almost membraneous laterally. Segment 12
heavily sclerotized throughout, having the same blackish colour as anterior three segments
dorsally. Posterior margins of segments 2, 3, and 10 dorsally with a row of slightly
sclerotized papillae, these membraneous only on venter. Anterior spiracle (Fig. 14)
with 9 tot 11 buds with usual combinations 9—9, 9—11, 10—11, 11—11, or 9—10.

The mature larva of pullitarsis (Plate 6c) is 6.5—9 mm long and 1.5—2 mm wide,
with sclerotization on segment 1 similar to that in rwfitarsis. Dorsum of segment 1
(Fig. 11) more or less heavily sclerotized, lateral area almost membraneous, and venter
evenly and slightly sclerotized, without any distinct pattern. Slight sclerotization also
visible on the anterior ridge. No rows of membraneous papillae have been found on
posterior margins, but distinct papillae are visible on anterior ridge and among the
striation on segment 1 ventrally; this segment distinctly striated both ventrally and
dorsally, segment 2 on anterior half dorsally and ventrally, segment 3 on anterior half
ventrally only. Otherwise the integument is smooth, except for regularly spaced, small,
flower-like papillae. Anterior spiracle (Fig. 15) with 4 to 5 buds, the number on each
side often differing.

The mature larva of rufttarsis (Plate 6d) has in general much in common with
pullitarsis; it is rather smaller, 6—8.5 mm long and 1.5—2.5 mm wide, more or less
sclerotized only on segment 1. The dorsal sclerotization (Fig. 12) is not very distinct
(light brown in colour), almost interrupted laterally near anterior spiracle and again
more distinct on a triangular patch ventrally. Papillae on posterior margins always mem-
braneous and therefore inconspicuous. Whole of segment 1 distinctly striated except
for the anterior ridge, which bears very small, not sclerotized papillae. Segment 2 with
fine striae on anterior half ventrally, otherwise segments smooth. Anterior spiracle
(Fig. 16) with 6 to 9 buds in combinations 7—8, 8—9, 7—7, 6—7, or 6—6.

The mature Lipara larvae may be keyed as follows:
1 Larva heavily sclerotized (blackish-brown in colour) on both tips, anteriorly on 4,
posteriorly on 2 segments. Anterior ons with 9 to 11 buds. Larva about 8 mm

[On ce A See rito
— Larva less AT oi oh in lo), ci antes at most on
the three anterior segments. Posterior segments pale, membraneous. . . 2

2 Three anterior segments slightly sclerotized on dorsum. All thoracic segments with
a row of heavily sclerotized papillae on posterior margin, segment 1 without a
ridge on anterior margin. Anterior spiracle with 11 to 12 (or rarely up to 14)
buds. Larva about 10 mm long and very stout. . . . PM acer’

-— Only segment 1 slightly sclerotized. Sclerotized papillae on ost margins of
thoracic segments absent, segment 1 with a distinct ridge on anterior margin.

Fig. 13—16. Anterior spiracle of mature larva of Lipara. 13, L. lucens Meig.; 14, L. similis Schin;
15, L. pullitarsis Dosk. et Chv.; 16; L. rufitarsis Loew. Scale = 0.1 mm

M. CHVALA, J. Dosko£ır.. I. H. Mook & V. POKORNY : The genus Lipara

10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

Anterior spiracle with at most 9 buds. Larvae generally smaller, about 8 mm long,
and ‘slender 1, EREN EE PSE ER ARR ER TAO
3 Anterior spiracle with 6 to 9 buds. Segment 1 usually slightly sclerotized, also
ventrally, distinctly striated on the whole surface, except the smooth anterior ridge;
segment 2 with slight striae on anterior half ventrally; segment 3 smooth, without
Straen etn PAN AS A EREN EEE I e E PN OE
— Anterior spiracle with 4 to 5 buds. Segment 1 usually membraneous beneath, at
most quite indistinctly sclerotized, but with distinct striae throughout, also on
anterior ridge; segment 2 with striae on anterior half both ventrally and dorsally;
segment 3 with striae on anterior half ventrally. . . . . . . . pallitarsis

Puparia

The puparia are brown to reddish-brown, with the same specific characters as in
mature larvae, i.e. of the same size, different colour of tips, and the same number of
buds on the anterior spiracle. L. /zcens has a large and stout puparium, 8 to 12 mm long,
the anterior two segments distinctly darkened dorsally, anterior spiracle with 11 to 14
buds; puparium of similis slightly smaller on average, distinctly blackish over three
anterior and two posterior segments, anterior spiracle with 9 to 11 buds; rufitarsis
and pullitarsis have smaller puparia, about 6 to 9 mm long, slightly darkened on
segment 1 only, anterior spiracle with 6 to 9 buds in r#fifarsis and 4 to 5 buds in
pullitarsis.

3. LIFE CYCLE AND INFLUENCE ON THE REED STEMS
a. Life cycle and gall formation

By their action in the larval stage, all four Lipara species induce the formation of a
terminal gall on the infested shoot, thus altering the length of the stem, the number
and dimensions of internodes and the formation of a panicle. Each gall is ultimately
inhabited by a single larva, even when shoots are infected by more than one egg.

The formation of the gall is principally the same in all four species, and is closely
related to the behaviour of the larvae. The egg-laying behaviour of the fly is described
in section 4. Several days after the eggs are laid the larvae hatch. In L. Iwcens the larvae
hatch after about 9 days. Hatching takes place virtually only during the night. This is
probably important for survival, because in order to escape drying out the larvae require
a very high humidity (Ruppolt, 1957, Mook, 1967). The young larva crawls upward
on the surface of the shoot untill it reaches the top part, and then crawls under the
edge of a leaf sheath into the roll of young leaves. There it feeds on the young leaf
parenchyma and bores downward toward the growing point. Just above this point it
continues to feed on the newly-formed leaves. Here the larva molts twice. The growing
point is affected by the feeding of the larva and probably also by its metabolic products.
Internodes formed subsequently remain shorter than the internodes of a healthy stem.

The shortening of the internodes leads to the formation of the terminal gall in the
stem. The gall itself is formed (except in the case of L.lwcens) by the leaves growing
from the agglomerated nodes. The leaf-sheaths do not have sufficient room to grow
parallel with the stem axis and are pushed off at an angle to it; in this way the typical
cigar- or spike-like inflation develops.

Up to this point the life cycle is basically the same for all the four species. The

M. CHVALA, J. DoskoCit, J. H. Mook & V. Pokorny : The genus Lipara 11

differences become apparent at a later stage, and therefore the rest of the life cycle will
be described for each species separately.

During July or August the larva of L. /ucens (Plate 6a) bites through the growing
point and lives in the shortened internodes where it feeds on the soft parenchyma filling
these internodes. In due time it gnaws a chamber passing into the 6th to 8th internode
from above. At the end of the summer and in the beginning of the autumn the larva
turns 180° in the chamber, so that its head is pointing upward. This turn is very
important (and takes place in the other species, too), because it enables the adult to
leave the gall. After turning, the larva shifts its position 2 to 3 cm higher, living without
nutrition approximately 1 to 2 cm under the remnants of the growing point; at that
time it is 8—12 mm long, thick, and ivory white. In this position it also hibernates
and pupates in the spring. In Czechoslovakia and in the Netherlands, depending on the
climatic conditions, the adults emerge as the second Lipara species at the end of May
or beginning of June (Fig. 17).

I
Q
D
Q

FEBR. MARCH APRIL MAY JUNE JAN. FEBR.
ERBUÜCENS ee L. RUFITARSIS
L.SIMILIS L. PULLITARSIS

Fig. 17. Life cycle of Lipara species in Czechoslovakia

The growing point of the reed is also perforated by the larva of L. rufitarsis (Plate
6d), but in this species the chamber reaches only to the 3rd or 4th (shortened) inter-
node from the top. The walls of the shortened internodes are not thickened and their
width is the same as that of the adjacent not shortened ones. Once arrived below the
growing point the larva feeds until some time around August, after which it turns and
rests until spring, when it pupates. At that time it is 6—8.5 cm long, slightly yellowish,
and more flattened than the larvae of the other species. The adult emerges as the
third Lipara species at the beginning of June.

12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

These two species are characterized by a common behaviour pattern viz., biting through
the growing point and life in the chamber in the shortened internodes. The other two
species leave the growing point intact, living throughout their whole lifespan between
the leaves above this.

In the beginning, the larva of L. pullitarsis (Plate 6c) feeds on young rolled leaves
in which it “chops” its chamber. Later on, when the panicle is beginning to be formed,
it feeds on this rudiment often completely destroying the panicle. Sometimes, probably
when infestation occurs later, a panicle large enough to survive is formed in the gall,
only to have the larva gnaw through it, too, to maintain its chamber. Such panicles never
develop fully. Before turning, the larva usually reaches the growing point, but after
having turned, it shifts itself 3 to 5 cm above the growing point and hibernates there.
At that time it is 6.5—9 mm long and white. In the spring, as soon as the days become
warmer, the larva pupates as the first of the Lipara species and the adults emerge also
as the first, in the second half of May.

Similarly, the larva of L. similis (Plate 6b) lives in the tube of leaf sheaths. In the
beginning it feeds on the rolled leaves, which later becomes completely severed from the
growing point, so that they can be pulled out by the tips. The space between the leaves
is relatively narrow and the adult larva fills it completely. As soon as the panicle begins
to develop the larva starts to feed on the rudiments, thus destroying the panicle com-
pletely. Even in the narrow cleft in the tube the larva turns and shifts itself upward.
During the winter and early spring, the two sheaths become hard, thus protecting the
hibernating larva, which at that time lies 5 to 7 cm above the remnants of the growing
point. In this period it reaches a length of 5.5 to 10 mm, and is slim, milky white, and
glossy, with both ends black. In the same position, the larva pupates in spring as the
last of the Lipara species. The adults emerge, also as the last, in the middle of June.

b. Description of the galls

The number of shortened internodes is more or less characteristic for the species and
thus also for the shape of the gall (Fig. 18 and Plate 4). The most massive and
prominent galls result from the influence of the larva of L. lucens (Plate 5a), and this
corresponds with the greatest number of shortened internodes, usually 10—13. The
shortened internodes are 2 to 3 times wider than the adjacent normal ones, and their
walls are thickened and strongly lignified (Plate 4a). All of the 6 to 8 uppermost
shortened internodes are brownish and covered with a continuous hairy layer. The
agglomerated nodes are also provided with a wreath of hairs, and the leaves growing
from them have short broad sheaths and weaker short blades. The highest leaves are
relatively broad and short, again covered with a continuous hairy layer; these lena do
not differentiate into sheath and blade.

Due to the activity of the larvae of L. rzfitarsis or L. pullitarsis, only 5 to 6 internodia
are manifestly shortened (Plate 4c). The galls (Plate 5c, d) in these two species
are usually similar, and can only be conclusively differentiated by a longitudinal section
through the gall. The galls are medium-sized to large, the pullitarsis galls being usually
larger, since pullitarsis occurs on wider shoots. This is why its galls are sometimes larger
than those of /zcens. But if the galls of lwcens and pullitarsis appear simultaneously on
equally wide shoots, those of lwcens are always larger (Fig. 20).

The stem infested by L. similis has the smallest number of shortened internodes, i.e.

PLAAT 1

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

*(AUIOYOG “A ‘30Yg) MaOT szsargufma

"I ‘p

“AUD 39 ‘YSOQ Szsartmnd “JT ‘D SUIS szprwis "TÈ

rie,

q “Bro Suasn] 7 PV vardi] JO py ‘I >Yeld

The genus Lipara

KORNY

M. CHVALA, J. Doskocit, J. H. Mook & V. Po

PLAAT 2

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

X 160 (Phot.

lis Egg: X 70, detail:

a UGC DSS @ chel,

’

il.

Plate 2. Lipara eggs with deta

J. Chalupsky)

M. CHVALA, J. Doskocit, J. H. Mook & V. POKORN

‘a

The genus Lipar

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 PLAAT 3

Plate 3. Lipara eggs with detail. a ‚b, L. pullitarsis; c, d, L. rufitarsis. Egg: X 70, detail X 160
(Phot. J. Chalupskÿ)

M. CuvAra, J. Doskoém, J. H. Mook & V. POKORNY : The genus Lipara

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 PLAAT 4

Plate 4. The highest internodes of reed stems with a gall of Lipara. a, L. lucens; b, L. similis;
c, L. pullitarsis (Phot. V. Pokorny)

M. CHVALA, J. DoskoCit, J. H. Mook & V. POKORNY : The genus Lipara

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 PLAAT 5
er

Plate 5. Galls typical of the individual species of Lipara. a, L. lucens; b, L. similis; c, L. pullı-
tarsis; d, L. rufitarsis. The size relations are maintened (Phot. V. Pokorny)

M. CHVALA, J. Doskocit, J. H. Mook & V. Pokorny: The genus Lipara

PLAAT 6

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

Yor”

d, L. rufitarsis (Phot. V.

L. pullitarsis;

)

b, L. similis;

rva of Lipara. a, L. lucens

Plate 6.

y)
J. H. Mook & V.

korn

The genus Lipara

POKORN

KOCIL,

ALA, J. DOS

ALA

. CHV

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 PLAAT 7

Plate 7. Signals of Lipara species registered with an UV recorder. a, signal of male L. /zcens with
part of answering signal of female; b-e signals of males of different species: b, L. /ucens; c, L.
similis; d, L. pullitarsis; e, L. rufitarsis

M. CHVALA, J. DoskoCit, J. H. Mook & V. POKORNY : The genus Lipara

M. CHVALA, J. Doskocit, J. H. Mook & V. Pokorny: The genus Lipara 13

AVERAGE LENGTH OF INTERNODES (CM)

4 2 3 4 5,6 TMD meten On 4e WAZ.
INNETNETRENNONDNESS

Fig. 18. Length of the highest internodes (counted from above) of reed stems attacked by larvae of
Lipara species. (average of 30 stems per species)

3 to 4. This is why the gall is the smallest and least noticeable (Plate 5b). Due to this
small number of shortened internodes and also to the fact that the shortened internodes
are narrower than the adjacent normal ones but rather telescopic (Plate 4b), the
sheaths have enough room and embrace the stem closely. The gall itself is formed only
by the leaf sheaths growing from the two highest visible nodes which form an approx-
imately 9 cm long tube concealing the remnants of the rolled leaves, the larva and the
remainders of the panicle and the growing point. This is why the gall is not inflated
and resembles the top of a sterile shoot, from which it differs only by the fact that
the sheaths of the two uppermost leaves covering the gall are almost equally long.
By summarizing these descriptions we arrive to the following key to the galls:
1 There is a terminal inflation on the stem: a cigar- or spike-shaped gall. . . 2
— There is no terminal inflation of the stem, it looks like the top of sterile shoot.
In the tube formed by the leaf sheaths growing from the two highest visible nodes
there is a slim, milky white, glossy larva with both ends black (or brown pupa with
both ends black) . . . MORT 0
2 Ina longitudinal section through the Di a Abe uo doni the young rolled

14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

leaves can be seen. In the chamber there is a big white larva (or brown pupa). The
chamber never passes SE the “ua B to the shortened internodes .

pullitarsis
— The lano n tai ie rest ci the growing point in order to reach the
shortened internodes. Its inner walls blak. . . . 3

3 Walls of the shortened internodes thickened, their tn two or Bin dim that
of adjacent not shortened ones. The chamber passing through 6 to 9 shortened
internodes, containing a big, thick, ivory white larva (or a brown pupa). The
shortened internodes brown . . . EN lacens

— Walls of shortened internodes neither Lidi nor bon their width equal to
that of the adjacent not shortened ones. The chamber passing through 2 tot 4
shortened internodes only and containing a big, yellowish, and somewhat dorsoven-
trally flattened larva (or brown pipa), A 4

4. BEHAVIOUR OF ADULTS

The discussion of behaviour and ecology will be somewhat one-sided, because Lipara
lucens is by far the best known species of the four and the outline of its biology will
therefore be the largest. The fragmentary information concerning the biology of the
other species will be compared with that of /ucens.

a. Precopulatory behaviour

The adults of Lipara lucens force themselves upward between the leaf sheaths and
emerge from tip of the gall during the early morning hours. In this process an important
role is played by the ptilinum, a large sac of the head, which can be expanded, so pushing
aside any obstacle in front of the head. It is also useful for bursting the puparium, and
in other species of higher flies, for forcing a way up through the soil (Oldroyd, 1964).
In the hardening process after emergence, the ptilinum is retracted inside the head.

The flies emerge over a period of two or three weeks in the end of May or the
beginning of June. Our observations do not indicate that the female flies emerge a few
days before the males, as is stated by Waitzbauer (1969).

The female flies are probably very sedentary before copulation, but detailed ob-
servations on the behaviour in the field are not yet available. In laboratory experiments,
where the flies are observed in small cages provided with some old reed stems, the
females remain in the same place for a long time unless the temperature becomes high
(above about 25° C), when they become restless. The males move about much more,
and it is often obvious that they are in search of a female. They fly from reed stem to
reed stem, on each stem giving a signal by vibrating. A virgin female fly sitting on the
same substrate as the male perceives this signal as a vibration of the substrate and im-
mediately answer by vibrating in its turn (Plate 7a). This in turn induces in the male
increased searching on the reed stem. It starts to walk over the stem, stops after a few
seconds to give a signal again, and when this is answered, begins to walk again. In this
way the male finally finds the female, and copulation often takes place without obvious
further display.

When a male gives its signal on a stem and does not receive an answer, it soon
leaves the stem and flies to another. Therefore the function of this behaviour seems
obvious. This procedure requires much less time than the male would need for searching
the whole reed stem.

M. CHVALA, J. Doskocit, J. H. Mook & V. Pokorny: The genus Lipara 15

The signals of both the male and the female fly cannot normally be heard by the
human ear, but the male’s signal is perceived when seated on a reverberating substrate.
Both signals can be made audible by placing a crystal gramophone pick-up element in
contact with the substrate, for instance a reed stem, and by connecting this element to a
tape recorder. The tape recordings can then be converted to visual signals with the help
of an oscilloscope or an UV recorder (Plate 7).

This description is based primarily on observations of L.lwcens (Mook and Brugge-
mann, 1968, 1971), but behaviour of other species is known to be similar. Important
differences between species were found in the signals of the males, characteristic examples
of which are given in Plate 7b-d. The main pitch of these sounds is about the same
(a few hundred Hz.), but there are wide differences in the rhythm making it easy to
recognize the different species by their signals. There are no great differences between
the signals of the females. They all have a pattern similar to the one shown in Plate 7a;
only the duration varies, that of L. pullitarsis being characteristically short (one or two
seconds) while that of the other species is usually longer (up to over 10 seconds). No
differences were found between the signals of flies collected in Czechoslovakia and the
Netherlands.

Because such marked differences were found between the signals of the males, an
experiment was set up in order to investigate whether these differences play a role in
reproductive isolation. In this experiment, sounds of three species of Lipara and two
species of Haplegis (small Chloropid flies, the larvae of which occur as inquilines in
Lipara galls) were played back to females of the three Lipara species. The female flies
rested on a strip of paper in contact with a small loudspeaker (output 0,5 watt). The
tape gave each of the signals ten times in a random sequence, with a 30 second interval
between signals. The tape was played back twice to each of five females of the three
different species, so that each female was subjected to each signal twenty times, and their
response was observed with a binocular microscope. The number of responses (Table I)
was high only for the own species, which makes it probable that the signals do indeed
play a role in reproductive isolation.

Table I. Reactions of 5 females of 3 Lipara species to 20 male signals of 5 different Chloropidae

3 signal Reactions of 9

lucens pullitarsis rufitarsis
L. lucens 18 16 18 19 18 — — — — — — — — — —
L. pullitarsis u Ze a ea EEE CL CERN
L. rufitarsis Zeelse mer. — — 19, 19720: 17 17
Haplegis spec. A SS SD ol, ai er
Haplegis spec. B - Jos oo ol oo do —

b. Egg-laying behaviour

The egg-laying behaviour of Lipara lucens has been investigated extensively (Mook,
1967). The species is autogenic: it does not need food to lay eggs. At emergence the
ovaries contain almost mature eggs, and egg-laying can start two or three days later.
The number of eggs in the ovaries of L. lucens lies between 64 and 96, with a mean
of 84 (data from 40 Dutch specimens). However, egg production is not limited to this

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

number. In laboratory experiments the flies were often observed laying eggs in two or
three periods of one or two days, each period separated by two or three days in which
no eggs were laid. The number of eggs laid during the first of these periods cor-
responded to the number of mature eggs in the ovaries at emergence. In the later
periods a smaller number of eggs was laid, so that the total production of eggs often
attained twice the number of eggs originally present in the ovaries. Under favourable
circumstances (the flies were provided daily with fresh drinking water and pieces of
fresh reed shoot of the preferred diameter; light and a temperature of 20° C were
maintened for 7 hours each day) the number of eggs laid was between O and 199, with
a mean of 133.6 (median 147).

Comparable data for the other species are lacking. At emergence they too have almost
mature eggs in the ovaries. For L. pullitarsis the number of eggs in the ovaries lies
between 98 and 157 in 20 Dutch specimens and for L. rufitarsis between 54 to 83 in 17
Dutch specimens. No data are available for L. similis.

Lipara lucens lays its eggs during the warmest hours of the day. The female flies to
a reed shoot and walks on it, first upward. After reaching the top of the shoot it often
turns and walks down again. During this walking the ovipositor may be slightly extended,
but does not touch the shoot. After a while the female begins to walk on a more restricted
part of the shoot, a few centimeters’ distance. By this time the ovipositor is fully extended
and its tip now and then touches the surface of the shoot. The fly then pauses for about
60 seconds with its head pointing upward, after which the egg emerges from the tip of
the ovipositor and is pressed against the surface of the shoot. After the egg has been
laid the female walks upwards and usually flies away to another shoot.

The eggs are usually laid on the stem, sometimes but not always on or under the
hairy ligules, but in an outdoor cage an appreciable proportion of the eggs was found
on the reed blades. The flies show a marked preference for ovipositing on shoots of a
certain diameter; nonpreferred shoots are generally deserted during the walking stage of
this ovipositional behaviour. Because this preference influences the habitat relationships,
it will be treated in the next section.

5. ECOLOGY
a. Habitat relations

In the literature it is often stated that the galls of Lipara lucens are most numerous
in reed stands located in dry places (Wagner, 1907; Docters van Leeuwen, 1957;
Ruppolt, 1957), but this does not necessarily imply that the water level itself influences
the species. Together with the water level, characteristics of the vegetation vary consider-
ably. The number of shoots per square metre, for instance, tends to be higher in drier
places (but is lower again where the soil becomes very dry). The dimensions of the
reed shoots are also influenced by the water level, the shoots being shorter and thinner
in dry than in wet places.

A partial regression analysis of samples of three different vegetations showed that
the variation in gall density is not primarily due to variations in water level or shoot
density, but to variations in the dimensions of the shoots. In this analysis and in the
subsequent experimental work the basal diameter of the shoots was chosen as the most
characteristic shoot dimension, because this is the most constant measure throughout the
year and is not influenced by gall formation. The basal diameter does not alter during

M. CHVALA, J. DoskoCit, J. H. Mook & V. PoKoRNY: The genus Lipara 17

the growing season and when the reed dries in the autumn it diminishes only by about
10 per cent.

Experimental work in the field and in the laboratory showed that the diameter of the
reed shoots plays an important role in the biology of Lipara lucens, in some cases directly
affecting the behaviour of the fly or its parasites, in other cases correlated only with the
real operating factor. The correlation of life functions of Lipara lucens with the diameter
of the reed shoots has been treated elswhere (Mook, 1967); only a short survey will
be given here.

The female flies lay their eggs preferentially on shoots with a width of 4 and 5 mm
at the base. In Fig. 19a oviposition is expressed as the mean number of eggs per shoot,
thus eliminating the influence of the number of shoots available in each class. Waitzbauer
(1969) found a distribution of eggs in the field with an optimum on shoots with a
diameter of 4—4.5 mm, but he did not take into account the distribution of shoot

0.6
04

0.2 d.

eggs per shoot

0.0 ovıposition preference

0.6

survival of larvae up
0.0 to gall formation

0.2

0.8
0.6

0.4
C.

fraction of surviving larvae

0.2
survival of larvae
0.0 in the gall
2 3 4 5 6 4 8 mm

basal shoot diameter

Fig. 19. Lipara lucens Meig. Correlation of oviposition and survival of larvae with diameter of
reed shoots

18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

diameter in the vegetation. He states that Mook (1967) found a preference for shoots
with a diameter of 2.5—2.9 mm, but this is derived from a laboratory experiment in
which the diameter of the shoot top was used. This value corresponds to a basal diameter
of about 4 mm.

Once a shoot has received an egg, the probability that a gall will be formed is also
correlated with shoot width (Fig. 19b). It is in the shoots with a diameter of about 2
and 3 mm that the young larvae have the greatest chance to induce a gall and thus the
greatest chance to survive. The differences in survival of larvae in the gall (fig. 19c)
are due to a heavier parasitism by Stenomalina and more predation by birds on the
thinner shoots.

Multiplication of the estimates of oviposition preference and survival up to gall

=}
©

diameter of the gall (mm)

6.0

20 30 40 50
diameter of the shoot base (mm)

M. CHvÂLA, J. Doskocit, J. H. Mook & V. POKORNY : The genus Lipara 19

formation in one diameter class gives the probability that a shoot in that class will develop
a gall. This appears to be highest on shoots with a diameter of 3 and 4 mm, and this
estimate agrees with the fractions of galled shoots in each class found in the field. In the
field a low number of galls is also found on shoots with a diameter of 7 or 8 mm and
sometimes even more, so that the chance of gall formation cannot be zero as in the
experiment of Fig. 19b. The experiments were of course, all carried out with a limited
number of shoots, so that chance variation can be important.

This analytical approach explains why the highest density of galls can be found in

E 100

diameter of the gall (m

6.0

4.0

20 30 40 50
diameter of the shoot base (mm)

Fig. 20—21. Correlation between basal diameter of reed shoot and diameter of gall occurring on

that shoot, for different types of galls: lignified galls containing larvae of L. /ucens (LJ) and

L. rufitarsis (WA), and soft walled galls with larvae of L. rufitarsis (©) and L. pullitarsis (9).

Galls collected in Voorsterbos (Fig. 20: parcel T 35, Fig. 21 parcel S 18), N.O. Polder, The
Netherlands 11-1-1963

20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974

places where a large proportion of the reed shoots have a width of about 3 or 4 mm.

It may be concluded that most galls of Lipara lucens are found in reed with sub-
optimal growth, but not in reed with the worst growth. Although the water level has
no direct influence, it can be said that reed with suboptimal growth is found mainly
in dry places. On the basis of experience with a wide variety of reed vegetations in the
Netherlands it can be said that there is no predilection for reed growing in extensive
vegetations, as compared to reed in smaller patches, for instance in small bogs, along
ditches, etc.

Lipara rufitarsis and especially L. pullitarsis are not as abundant in the Netherlands
as L. lucens. Only superficial observations on their habitat are available, but comparison
with L. lacens is possible. In reed vegetations where two or three species are present the
galls of L. rufitarsis and L. lucens occur on shoots with about the same diameter, but th
shoots with galls of L. pullitarsis on the average have a larger diameter. This can be seen,
for instance, in Fig. 20, but has also been found in many other vegetations.

Galls of L. rufitarsis are sometimes present in low numbers in closed reed vegetations
on dry soil, together with L. lwcens, but occur much more abundantly in smaller reed
vegetations in dry places at the edge of woods or along roadsides or other somewhat
shaded places. In such places they are often more numerous than L. lucens.

Galls of L. pullitarsis were found in small reed stands with a low shoot density (often
located between trees or bushes) in wetter, marshier places than those where L. rufitarsis
has its optimum. In the Netherlands no galls of pullitarsis were found in large closed
reed vegetations. This shows clearly that shoot dimension is not the only important factor
determining the habitat of Lipara species.

In places where L. rufitarsis and L.lucens occur together, the larvae of the former
species are sometimes found in galls that are indistinguishable from those of the latter
species. This peculiar phenomenon has already been reported by Ruppolt (1957), but
he did not describe L. rufitarsis in its typical galls. The thin galls with larvae he
describes under the name of L. rufitarsis, undoubtedly refer to L. pullitarsis.

The two types of galls in which rwfitarsis is found are quite different. That no inter-
mediate forms occur is evident, e.g. in Figs. 20 and 21, showing two distinct groups
of dots. The upper group corresponds with the wide galls with thickened internodes and
lignified walls, the lower group, with narrow galls of which the internodes are not
thickened. For both types of gall there is a strong positive correlation between shoot
and gall width illustrating that the diameter and other dimensions of a gall are significant
only in relation to shoot diameter.

Analysis of the morphology of the larvae and adults and also of the acoustical signals
of the male flies, proved that the same species (rufitarsis) occurs in both types of gall.
A few galls of the /ucens type have also been found in which only a pullitarsis larva
was present (above the growing point).

It is difficult to believe that one species can induce different galls on the same food
plant. We know that when a shoot is infected by more than one egg, never more
than one larva will survive. When these eggs are of different species, it cannot be said
beforehand which species will survive. In the case of multiple infection by /zcens and by
one of the smaller species, if the /ucens larva dies the result will be a typical /ucens gall
with a surviving rufitarsis or pullitarsis larva. This only holds, however, if the /ucens
larva does not die before the gall reaches maturity, since otherwise we would find a gall
of intermediate size. We know that gall growth stops when a /zcens larva dies before

M. CHVALA, J. Doskocit, J. H. Mook & V. POKORNY : The genus Lipara 21

gall maturation, because we find rudiments of galls above which the shoot has resumed
normal growth. This does not imply, however, that /zcens will always succumb to the
smaller species. Without extensive breeding experiments, it will be impossible to judge
the outcome of interspecific competition.

b. Parasites

Hymenopterous parasites belonging to four families attack the different species of
Lipara. We have not tried to give a comprehensive list of these parasites. Only the
common species with which we are acquainted will be treated here, because the records
of occasional parasites are difficult to check. It is important to rear from material found
in opened galls, to be sure that the parasites originate from Lipara larvae and not from
the numerous species of inquilines that are also present in the galls.

The occurence of parasites differs widely between the types of galls. Table II shows
the numbers of hosts and parasites found in galls collected in a number of localities in
the central and eastern part of the Netherlands. The /zcens galls included here originate
exclusively from places where this species occurred together with one or both of the
other fly species. More figures on parasitism of L./ucens have been given elswhere
(Mook, 1967).

The findings of Table II cannot be generalized for other regions. The relative im-
portance of the parasites in different regions is indicated below under the different
species.

Table II. Distribution of host and parasite larvae over 3 types of galls collected in various places
in the Netherlands

Type of gall

Species of larvae lucens rufitarsis pullitarsis
Lipara lucens 191 — —
Lipara rufitarsis 31 240 —
Lipara pullitarsis 9 — 119
Polemon liparae 83 me ae
Stenomalina liparae 58 8 13
Scambus phragmitidis 1 1113 4
Total 373 361 136

Ichneumonidae: From Table II it is apparent that Scambus phragmitidis Perk. is
primarily a parasite of L. rufitarsis. Waitzbauer (1969) reared one specimen from /ucens
in Austria (Neusiedler See). In the Netherlands we have found a few in galls of /ucens,
but always in places where rufitarsis also occurred in /ucens galls. This parasite was
erroneously named Pimpla arundinator F. var. similis Bridgm. in Mook (1967). The
new identification was made by Mr. K. W. R. Zwart, Wageningen, who compared
specimens with paratypes from the British Museum (Natural History), London.

The distribution is difficult to assess. Pokorny did not find Scambus in Czechoslovakia.
Other species of the same genus have been mentioned in the literature: Giraud (1863)
reared Scambus (= Pimpla) arundinator F. as a parasite of L. rufitarsis or L. pullitarsis
in Austria, while Wagner (1907) in Germany (Hamburg) and Blair (1944b) in England
mention Scambus (= Pimpla) detritus Holmgr. from galls of L. /ucens.

22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
Cnemopogon apicalis
5
demen
10. Scambus phragmitidis
| À
0 ERK

is Lipara pullitarsis

Polemon liparae

number of specimens
Ui oO

(©)

= Lipara lucens
x PA

Lipara rufitarsis

Stenomalina liparae

5 chr i li

24 26 28 30 2 4 6 8 10 12 14 16 18 20 22 24 26 28
APRIL ~ MAY

Fig. 22. Emergence in the laboratory of adult flies, predators and parasites from galls collected in
N.O. Polder and near Dalfsen, The Netherlands, 20-IV-1962

M. CHVALA, J. DoskoCr, J. H. Mook & V. Pokorny: The genus Lipara 23

Of the life history of Scambus phragmitidis nothing more is known than that it
emerges early in the season (see Fig. 22), long before L. rufitarsis. The oviposition
behaviour has not been studied.

Braconidae: Two species of Polemon have been described: P. liparae Gir. and P. melas
Gir. In the Netherlands P. liparae is primarily a parasite of L. lucens. In Austria (Waitz-
bauer, 1969) and Czechoslovakia (Pokorny) it is by far the most important parasite of
L. lucens, while in Hungary it was the only parasite found in this host (Erdös, 1961).
Giraud (1863) mentions this species as the only parasite of Lipara similis.

Polemon melas is the most important parasite of L.rufitarsis in Czechoslovakia
(Pokorny). Giraud (1863) found it in Austria. It was reared occasionally in the Nether-
lands (Reynvaan and Docters van Leeuwen, 1906).

Polemon liparae emerges from the tip of the gall just as its host and at the same time
as L. lucens. It oviposits in the eggs of /ucens (Mook, 1961). Larvae of lwcens parasitized
by Polemon form a puparium in September instead of in the spring, as they normally
do. A puparium containing a parasite can be distinguished from the normal by being
reddish-brown and slightly more slender. Varley and Butler (1933, see also Askew,
1971) have suggested that the premature pupation might be caused by wounding by the
ovipositing parasite. This is improbable because of the long time lying between infection
and pupation.

Pteromalidae. Stenomalina (= Pteromalus) liparae Gir. was found in the Netherlands
in galls of lucens, rufitarsis and pullitarsis (see Table II), but most commonly in /zcens.
In England (Blair, 1932) and western Germany (Hamburg, Wagner, 1907) it is the
most common parasite of /wcens. In Czechoslovakia it is not common (Pokorny); it was
not found in eastern Austria (Neusiedler See) by Waitzbauer (1969) and in Hungary
by Erdös (1961). Giraud (1863) has described it from material collected near Vienna
and indicates that it was rather common there.

The adults of Stenomalina liparae leave the gall by biting a small round hole in the
leaf sheaths just above the gall chamber. They begin to emerge about 8—10 days later
than L. lucens. They parasitize the young larvae living above the growing point, by boring
through the leaf sheaths. In view of the circumstance that the first larvae of /zcens arrive
above the growing point in the period that Stexomalina starts to emerge and that this
species does not need much time between emergence and oviposition, it seems that its
life cycle is primarily synchronized to that of L. /ucens.

Eulophidae. Tetrastichus legionarius Gir. is mentioned as a parasite of L. /ucens in
Czechoslovakia (Pokorny), Austria (Giraud, 1863), and the DDR (Greifswald, Ruppolt,
1957), but seems nowhere to be common. Erdös (1961) mentions the species from
Hungary as a parasite of Haplegis flavitarsis Mg., a small chloropid fly that is an in-
quiline in Lipara galls. In the Netherlands 7. legionarius is rare in lucens. We have
found only recently two parasitized puparia, after having opened more than 10.000 galls
from various places, although mainly from the eastern part of the country. Mr. M. J.
Gijswijt ('s-Graveland), who kindly identified our specimen, informed us that the species
has also been reared from /wcens galls from three localities in the central and southern
parts of the Netherlands. It is a gregarious parasite: our puparia contained 25 and 28
larvae respectively.

24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
c. Predators

Birds attack the galls of L.r#fitarsis in particular and also the smaller galls of
L. lucens occurring on narrow shoots and, less often, the galls of L. pullitarsis. The
intensity of this predation differs widely between winters. Blue Tits (Parus caeruleus L.)
are known to predate in winter on all kinds of insects in reed stems (Tischler, 1943).
Actual observations of predating tits are scarce, but they are at least known to be able
to open the galls (Mook, 1967). Kramer (1917) observed a Great Spotted Woodpecker
(Dryobatus major L.), opening a gall of L. lucens.

Puparia of Cnemopogon apicalis Wiedemann (Diptera, Scatophagidae) are found in
the winter in small numbers between the top blades of all three types of galls. The
host larvae are always absent, presumably having been consumed by the Cremopogon
larvae (Wagner, 1907; Theowald, 1961).

6. AKNOWLEDGEMENTS

We are indebted to Miss C. G. Bruggemann (Arnhem) for doing much of the work
on the acoustical behaviour, to Dr. H. PeSkova (Praha) for her generous help in the
investigation of the immature stages, and to Mrs. J. A. van Noort and Mr. D. Landsman
(Arnhem) for assistance in collecting data for sections 4 and 5. Thanks are also due to
Dr. J. Chalupsky (Praha) for making the photographs of Plates 2 and 3, to Prof. Dr.
L. M. Schoonhoven (Wageningen) for making the UV recordings of Plate 7 and to
Mr. W.J. N. M. Verholt for drawing Fig. 19—22.

7. REFERENCES

Askew, R. R., 1971, Parasitic insects. Heinemann, London, 316 pp.

Balachowsky, A. et L. Mesnil, 1935, Les insectes nuisibles aux Plantes cultivées. Vol. 1, Paris, 1137 pp.

Becker, T., 1910, Chloropidae. Eine monographische Studie I. Arch. Zool. 10: 33—174.

Blair, K. G., 1932, Some notes on the galls of Lipara lucens. Entomologist’s mon. Mag. London.

68: 10—13.

, 1944a, Further notes on the galls of Lipara lucens Mg. Entomologist’s mon. Mag. 80: 6—7.

, 1944b, Some galls of Lipara lucens Mg. Entomologist's mon. Mag. 80: 189—190.

Collin, J. E., 1946, The British Genera and Species of Oscinellinae (Diptera, Chloropidae). Trans.
R. Ent. Soc. Lond., 97: 117—148.

Docters van Leeuwen, W. M., 1957, Gallenboek, 2 nd. ed. Zutphen: Thieme. 332 pp.

Doskoäil, J. et Chvála, M., 1971, A revision of Lipara Meigen (Diptera, Chloropidae), including
the description of a new species from Europe. Acta ent. bohemoslov., 68: 100—107.

Duda, O., 1933, Chloropidae, in: E. Lindner Ed., Die Fliegen der palaearktischen Region. Stuttgart.
248 pp.

Enderlein, G., 1913, Analytische Tabelle der Gattungen der Chloropiden-Subfamilie Oscinellinae.

Zool. Anz. 38: 10—13.

, 1961, Beobachtungen über die Insektencönose des Schilfes. 8. Wanderversammlung

Deutscher Entomologen, 171—177.

Giraud, J., 1863, Mémoire sur les insectes qui vivent sur le Roseau Commun, Phragmites communis
Trin. et plus spécialement sur ceux de l'ordre des hyménoptéres. Verh. zool. bot. Ges.
Wien 13: 1251—1288.

Heeger, E., 1856, Neue Metamorphosen einiger Dipteren, Sber. K. Akad. Wiss. Wien. 20: 335—
345.

Hennig, W., 1952, Die Larvenformen der Dipteren. 3 Teil, Berlin: Akad.-Verlag, 628 pp.

Erdos, J.

M. CiivALa, J. Doskocit, J. H. Mook & V. POKORNY : The genus Lipara 25

Kramer, H., 1917, Die Musciden der Oberlausitz. Abh. Naturforsch. Ges. Görlitz. 28: 334— 338.
Loew H., 1858, Zwanzig neue Diptern. Wien. Ent. Monatschr., 2: 57—62.
De Meijere, J. C. H., 1949, Achtste supplement op de nieuwe naamlijst van Nederlandse diptera
van 1898. Tijdschr. Ent. 92: 1—14.
Mook, J. H., 1961, Observations on the oviposition behaviour of Polemon liparae Gir. (Hym.,
Braconidae). Archs. néerl. Zool. 14: 423—430.
— , 1967, Habitat selection by Lipara lucens Mg. (Diptera, Chloropidae) and its survival value.
Archs. néerl. Zool. 17: 469—549.

, and C. G. Bruggemann, 1968, Acoustical communication by Lipara lucens (Diptera, Chlo-
ropidae). Entomologia exp. appl. 11: 397—402.

, and C. G. Bruggemann, 1971, Acoustical behaviour of Chloropid flies living on reed
(Phragmites communis Trin.). Proceedings XIII int. Congress of Entomology, I; 418—419.

Oldroyd, H., 1964, The natural history of flies. Weidenfeld and Nicholson, London.

Pokorny, V., 1970a, Bionomie druhu Lipara lucens Meig. a L. similis Schin. na räkosu. Ziva, Praha,
3: 101-103.

—., 1970b, The influencing of reed (Phragmites communis Trin.) by gall forming Diptera of
the genus Lipara Meig. PT - PP Report No. 1 (1964—1969) Productivity of terrestrial
ecosystems, Praha, 135—136.

——, 1971, Flies of the genus Lipara Meigen on Common Reed, Hidrobiologia, Bucarest, 12:

287—292.
, 1972, Bionomie druhu Lipara rufitarsis Loew. a L. pullitarsis Dosk. et Chvála na rákosu.
Ziva, Praha (in press).

Reynvaan, J. and W. M. Docters van Leeuwen, 1906, Die Entwicklung der Galle von Lipara lucens.
Recl. Trav. Bot. néerl. 12: 235--261.

Ruppolt, W., 1956, Uber die cecidogene Diptere Lipara lucens Meigen (Chloropidae). Naturwissen-
schaften, 43: 260.

—., 1957, Zur Biologie der Cecidogenen Diptere Lipara lucens (Chloropidae), Wiss. Z. Ernst
Moritz Arndt-Univ. Greifswald, Mat.-Naturwiss. Reihe, 6: 280—292.

Sabrosky, C. W., 1941, An Annotated List of Genotypes of the Chloropidae of the World. Ann.

ent. Soc. Am., 34: 735—765.
, 1958, A Phragmites Gall-maker new to North America (Diptera, Chloropidae). Proc. ent.
Soc. Wash., 60: 231.

Schiner, J. R., 1854, Dipterologische Fragmente IV. Verh. zool.-bot. Ges. Wien 4: 169—176.

Theowald, Br., 1961, Diptera uit de sigaargal van het riet. Ent. Ber., Amst. 21: 108—109.

Tischler, F., 1943, Schilfrohr als Nahrungsquelle für insekten-fressende Vögel im Winter. Vogelzug.
14: 69—71.

Varley, G. C. and C. G. Butler, 1933, The acceleration of development in insects by parasitism.
Parasitology, 25: 263—268.

Vimmer, A., 1925, Larvy a kukly dvoukfidlého hmyzu stfedoevropsk&ho. Praha: Ceská graf. Unie,
348 pp.

Wagner, W., 1907, Uber die Galle der Lipara lucens Meig. Verh. Ver. naturw. Unterh. Hamb. 13:
120—135.

Waitzbauer, W., 1969 Lebensweise und Produktionsbiologie der Schilfgallenfliege Lipara lucens
Mg. (Diptera, Chloropidae). Sber. dst. Akad. Wiss. Mathem.-Naturw. Kl., Abt. I, 178:
175—242.

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PYRALIDAE OF THE THIRD ARCHBOLD EXPEDITION.
1. SUBFAMILY ODONTIINAE
(LEPIDOPTERA)

BY
EUGENE MUNROE
Ottawa, Canada*)

ABSTRACT

Nine species in six genera are recorded from the material of the Third Archbold Expedition
to Western New Guinea and are figured. The following taxa are described as new: Trigo-
noorda gen. nov., type-species T. triangularis spec. nov. also includiing T. iebelealis spec. nov;
Pseudonoorda gen. nov., type-species P. minor spec. nov., also including P. brunneiflava spec. nov.
and the previously described P. nigropunctalis (Hampson) comb. nov., P. distigmalis (Hampson)
comb. nov. and P. metalloma (Lower) comb. nov., ex Noorda, and P. noordimimalis (Hampson)
comb. nov., P. brunneifusalis (Hampson) comb. nov, P. photina (Tams) comb. nov. and P. lampra
(Tams) comb. nov., ex Clupeosoma; Hyalinarcha gen. nov., type-species H. hyalina (Hampson)
comb. nov., ex Boeotarcha. Thesaurica notodontalis (Hampson) comb. nov. and T. accensalis
(Swinhoe) comb. nov. are transferred from Sameodes and Noorda, respectively.

INTRODUCTION

The rich material of Pyralidae collected by the Netherlands Indian — American Exped-
ition to the interior of Dutch New Guinea (now West Irian), better known as the Third
Archbold Expedition, has been lent to me for study for a number of years, thanks to the
generosity and sorely tried patience of the Rijksmuseum van Natuurlijke Historie, Leiden.
During this time the whole of this material has been prepared and sorted to species and
the problems of identity and relationship have for the most part been solved. I now
propose to give lists of the material, subfamily by subfamily, with remarks on relation-
ships and with descriptions of new taxa where required. A general faunistic analysis
will be deferred until after the taxonomic treatment is complete, but the following state-
ments will serve for introduction.

The collection was made mostly in relatively undisturbed habitats at a variety of
elevations from near sea level to 3800 m. The localities are described in detail by Brass
(1941), Archbold, Rand & Brass (1942) and Toxopeus (1940). The most intensive
collecting was done at middle and higher altitudes, from which the species represen-
tation is good. Many species were collected at lower levels also, but they probably
represent a considerably smaller fraction of the fauna there. The fauna shows pronounced
altitudinal zonation. In a general way this corresponds well to what is known from
eastern New Guinea. The same groups: Scopariinae, certain Nymphulinae, etc., are well
developed at high altitudes, with such groups as Epipaschiinae and certain Phycitinae
and Pyraustinae becoming important in the 1500—2500 m zone. At both levels some
species are known from eastern New Guinea, whereas others are known only from the

*) Biosystematics Research Institute, Canada Department of Agriculture, Central Experimental Farm,
Ottawa, Ontario KIA OC6, Canada.
27

28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974

present collections. At the lower levels, especially Araucaria Camp at 800 m, at which
a large collection was made, the fauna is generally similar to that in other parts of
New Guinea.

The high-altitude material is the most extensive so far collected in New Guinea, but
the whole collection is of very great interest. In the main it reflects the hard and careful
work of the late L. J. Toxopeus, whose untimely death in 1951 interrupted a brilliant
entomological career.

This first paper deals with the subfamily Odontiinae. Although the subfamily has a
number of genera and species in New Guinea, the representation in the present collection
is poor. I can only speculate as to the reasons, but there are probably relatively few species
at high altitudes, while at moderate and low altitudes they may have been eclipsed by
the many more conspicuous and exciting species of other groups.

SUBFAMILY ODONTIINAE

Clupeosoma Snellen
Clupeosoma Snellen, 1880: 203.

Type-species C. pellucidalis [recte, pellucidale} Snellen, 1880, Celebes, by monotypy.

This genus was used by Hampson, 1897: 216, for a rather mixed assemblage of
odontiine and pyraustine species, which he placed in the Hydrocampinae (= Nymphu-
linae) because of the stalking of R, with Rz 4 4. In my “Synopsis of the North American
Odontiinae” (Munroe, 1961: 5) I referred Clupeosoma to the Odontiinae without dis-
cussion. As will be seen from the figure, the genitalia are typically odontiine, with
bilobed setose uncus, with gnathos arms fused basally with tegumen, with strong coremata
associated with the juxta, and with rounded, radially fluted valves.

I restrict the genus to a series of species closely related to C. pellucidale, including
C. astrigale Hampson, emend., from Queensland; C. subrufale (Rothschild), emend.,
comb. nov., from Ceram, described in Noorda; C. atristriatum Hampson, emend., from
Rookfe] Island; C. rufistriatum Hampson, emend., from the Oetakwa River, Snow
Mountains, New Guinea; C. glaucinale Hampson, emend., from Sumbawa; and C. la-
niferale Hampson, emend., from St. Aignan, as well as a number of undescribed species.
Snellen’s characterization of the genus is good, and will serve to distinguish it from the
discordant elements placed in it by Hampson. The maculation is characteristic and is
shared only by the closely related genus Neogenesis Hampson, discussed below.

Clupeosoma rufistriatum Hampson
Figs. 1, 10
Clupeosoma rufistriatum {sic} Hampson, 1917: 277.
The single specimen of the genus in the present collection appears to belong to

Hampson’s species, described from the Oetakwa River at 3.000 ft [ca. 900 m].
1g, Araucaria Camp, 800 m, 24 March 1939, L. J. Toxopeus.

Neogenesis Hampson

Neogenesis Hampson, 1907: 19.
Type-species N. flaviplagialis Hampson, from New Guinea, by monotypy.
As Hampson noted, this genus has clearly arisen from Clupeosoma by exaggeration

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 29

of the secondary sexual characters and by stalking and fusion of veins in both sexes.
The single known species is widely distributed in New Guinea.

Neogenesis flaviplagialis Hampson
Figs. 2, 11, 19

Neogenesis flaviplagialis Hampson, 1907: 19.

The species should not be mistaken for anything else. It was described from material
from Milne Bay (type locality) and Moroka, British New Guinea [now Papua}. Meek
collected it in the Snow Mountains and George Holland and I found it at several
localities in eastern New Guinea. I can see no significant geographical variation.

1g, Hollandia, 1938, Brinkmann; 14, Araucaria Camp, 800 m, 22 March 1939,
L. J. Toxopeus; 1g°, Mist Camp, 1800 m, 15 Jan. 1939, L. J. Toxopeus.

Trigonoorda gen. nov.

Type-species Trigonoorda triangularis spec. nov.

External characters. Frons slightly prominent, either rounded or flat, smoothly scaled.
Vertex short, with rough, erect scaling. Labial palpus porrect, exceeding frons by about
dength of head, scaling flattened, blade-like, third segment largely hidden in scaling of
second. Maxillary palpus prominent, with an expanded distal tuft of scales. Proboscis well
developed, scaled at base. Eye large. Ocellus present but small. Antenna filiform, in male
with feathery ciliations, several times as long as diameter of shaft; in female with short
even ciliations; in both sexes smoothly scaled dorsally. Body short but not especially
robust. Praecinctorium prominent, with transversely rounded distal tuft. Legs slender,
without unusual specializations.

Forewing broadly triangular; costa and termen somewhat convex; apex acute; posterior
margin sub-basally convex and with a weak scale tuft. Cell rather narrow, a, little over
half of length of wing. R, from cell basad of anterior angle. R, stalked with R3 + 4,
the latter long-stalked. R, from anterior angle of cell, weakly curved and approximated
to R3 + 4 at base. Discocellular straight, somewhat oblique. M, from somewhat behind
anterior angle of cell, diverging from R;. M, and M, from posterior angle of cell, their
origins close but slightly separated, their basal parts not or scarcely curved and approx-
mated. Cu, from well basad of posterior angle. Cu, from cell at two-thirds to three-fifths
from base. Anal loop large, very weak distally.

Hind wing of moderate width, termen rounded. Sc and R, anastomosed. R, and M,
stalked for some distance. Cell about half length of wing. Discocellular concave, posterior
limb strongly oblique distad. M, and M, from posterior angle of cell, their basal parts
usually weakly curved and approximated. Cu, from cell at two-thirds to three-fifths.
Cubital and anal area with slender, weakly spatulate, hairlike scales.

Male genitalia. Uncus rather narrowly subtriangular, tip rounded, sides rolled down
and weakly setose. Gnathos Y-shaped. Transtilla narrow, arched. Juxta large and
pyriform. Vinculum ventrally rounded and expanded, bearing moderately developed
submedial and lateral paired coremata. Valve of moderate width, ovoidal, with a sinuate
ridge well below costa and with inflated sacculus; distal and costal parts radially fluted;
a variably developed group of short spines on ventrodistal curvature at end of sacculus.
Penis slender and curved, with fairly long basal caecum; armature of vesica weak or
absent.

30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974

Female genitalia. Ovipositor with high narrow lobes bearing relatively few, variably
developed setae. Apophyses moderate to long. Eighth tergite rectangular or with anterior
margin medially produced, with scattered setae. Ductus bursae membranous, evenly
narrow, of moderate length, with a sclerotized collar at junction with bursa. Bursa round
or pyriform, membranous, with a small signum near junction of ductus bursae.

Remarks. This genus would key to Zebronia Hübner in Hampson’s Nymphulinae,
but it has only certain superficial characters in common with that pyraustine genus. It
is related in a general way to Clupeosoma and Autocharis. It differs from the former in
the shorter palpi, broadly triangular wings and in characters of the wing venation,
maculation and genitalia. From the latter it differs, inter alia, in the stalking of Ro with
Rg + 4.

In addition to the two species described here there are several smaller undescribed
species from New Guinea and the Sunda Archipelago, which will be dealt with else-
where.

Trigonoorda triangularis spec. nov.
Figs. 3, 12, 20

External characters. Frons somewhat rounded, dark reddish brown. Vertex contrastingly
light buff. Maxillary palpus reddish brown. Basal scaling of proboscis buff. Eye and
ocellus brownish fuscous. Antenna buff; dorsal scaling with fuscous segmental bands.
Thorax above anteriorly reddish brown, posteriorly light buff. Abdomen above buff.
Body beneath light buff. Legs whitish buff, with femora reddish brown and outer
surfaces of front tibiae and tarsi broadly banded with reddish brown.

Forewing above somewhat thinly scaled, light, weakly pinkish buff with fine fulvous-
buff dusting. Costa and termen narrowly dark reddish brown. Antemedial line narrow
weak, almost straight; oblique distad from costa at one-third from base to posterior
margin at two-fifths. Discocellular bar fine, fuscous, connected anteriorly with costal
line. Postmedial line narrow, weak, slightly dentate, tending to be interrupted at veins,
dark reddish brown or brownish fuscous; oblique distad from costa at four-fifths from
base as far as middle of cell Mg, there angled and curved increasingly basad to just in
front of Cuz, there obtusely angled and erect to posterior margin at three-fourths from
base. Fringe light buff basally, white distally.

Hind wing above translucent pale yellowish buff. Fringe whitish buff.

Forewing beneath light pinkish buff, with silky lustre. Costal line as above. Terminal
line more diffuse. Discocellular bar strong. Transverse lines visible only by transparency.
Fringe as above.

Hind wing beneath as above.

Expanse 21 to 24 mm.

Male genitalia. Uncus less than three times as long as wide. Gnathos with median
element tapering to a point distally. Dorsal part of juxta narrow. Patch of spines at
ventrodistal curvature of valve strong.

Female genitalia. Ovipositor lobe high and narrow, with long setae. Apophyses short.
Eighth tergite rectangular, with few fine setae, most numerous posterolaterally. Ostium
membranous. Ductus bursae short but slender and of even width. Bursa round, signum
near junction of ductus bursae.

Life history. Unknown.

Types. Holotype, g', allotype, 9, and 4 paratypes, Moss Forest Camp, 2600—2800

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 31

m, 11—24 Oct. 1939, L. J. Toxopeus. Holotype, allotype, and two paratypes in Leiden
Museum; one male and one female paratype in Canadian National Collection, Type
No. 12.861.

Remarks. This species closely resembles T. iebelealis but is slightly smaller, with fore-
wings sharper and less warmly coloured and with finer, sharper markings; the hindwings
are paler and less strongly yellowish in the present species. The genitalia differ in the
two species in a number of characters as noted in the descriptions.

Trigonoorda iebelealis spec. nov.
Figs. 4, 13, 21

External characters. Frons flat and oblique, dark brown medially, lighter towards sides
but with a dark border. Vertex reddish brown anteriorly, yellowish buff posteriorly. A
whitish-buff line between eye and antenna. Labial palpus with mixed fuscous and fer-
rugineous scales, its base beneath narrowly but contrastingly white. Maxillary palpus
fuscous; base of proboscis buff. Antenna light buff with segmental fuscous bands
dorsally. Thorax above reddish brown anteriorly, shading to warm buff posteriorly.
Abdomen above light greyish buff. Body beneath and legs whitish buff; outer surface
of front legs broadly banded with purplish fuscous.

Forewing broadly triangular; costa and termen somewhat convex; apex subrectangular;
termen slightly oblique, tornus rounded. Ground colour above warm yellowish buff,
finely dusted with ferrugineous. Costa ferrugineous. Antemedial line weak, narrow,
somewhat sinuated, ferrugineous; oblique distad from costa at one-third from base to
posterior margin at three-fifths. A weak fuscous discocellular bar.

Postmedial line slightly oblique distad from costa at five-sixths to vein Mg, there
weakly retracted to an obtuse angulation on Cu, and erect to termen near tornus. Termen
with a narrow, somewhat diffuse, ferrugineous band. Fringe yellowish at base, whitish
distally.

Hind wing above translucent yellowish buff, a little darker terminally. Faint traces
of a reddish-brown postmedial line in cubito-anal area. Fringe as on forewing.

Wings beneath much as above. Forewing a little darker, with transverse lines lost and
other dark markings reduced. Hind wing as above.

Expanse 23 to 25 mm.

Male genitalia. Uncus more than three times as long as wide. Gnathos with median
element finger-like and not tapering. Dorsal part of juxta wider than in T. triangularis.
Patch of spines at ventrodistal curvature of valve very weak.

Female genitalia. Ovipositor lobe high and narrow, but somewhat reduced and with
few setae. Apophyses long and strong. Eighth tergite elongate and heavily sclerotized,
produced forward medially and at anterolateral angles, its surface with scattered, fairly
strong setae. Ductus bursae wider than in T. triangularis, pyriform and with signum far
from junction of ductus bursae.

Life history. Unknown.

Types. Holotype, &, allotype, 9, and 1 9 paratype, TehelE Camp, 2250 m, 7 and
26 Nov. 1938, L. J. Toxopeus. 1 @ paratype, Sigi Camp, 1500 m, 23 Feb. 1939,
L. J. Toxopeus. Holotype, allotype and one paratype in Leiden Museum; one paratype
in Canadian National Collection, Type No. 12.862.

Remarks. This species is very similar in appearance to T. triangularis, but is a little

32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974

larger; the forewing is more orange and the hind wing is yellower; the transverse lines
of the forewing are a little more diffuse; the antemedial line is more oblique and less
regular. There are obvious differences in the male genitalia, especially the much weaker
development in the present species of the spinose patch on the ventrodistal curvature
of the valve.

Pseudonoorda gen. nov.

Type-species: Pseudonoorda minor spec. nov.

External characters. Frons flat and oblique, smoothly scaled. Vertex with rather smooth
vestiture of raised and often broad scales, arising laterally and curving anteromesad.
Labial palpus long and porrect, with flattened, blade-like, triangular scaling; third seg-
ment wholly or partly hidden in scaling of second. Maxillary palpus prominent, with
triangular distal scale-tuft. Proboscis well developed, scaled at base. Eye large. Ocellus
present. Antenna filiform, ventrally short-pilose in both sexes, dorsally scaled. Body robust,
abdomen slightly exceeding anal angle of hind wing. Praecinctorium with transverse,
rounded, distal fan of scales. Legs slender, outer tibial spurs somewhat shorter than inner.

Forewing triangular, costa and termen weakly convex, posterior margin almost straight.
Cell about half as long as wing, of moderate width. Discocellular erect anteriorly, curved
increasingly distad, posterior angle of cell moderately acute. R, from near end of cell,
closely apposed to Ro 4. Ry stalked for a considerable distance with R; + 4. Rz and
R, stalked to near apex. R; from anterior angle of cell, its basal part curved and approx-
imated to Rs 4 4. M; from anterior angle of cell, but straight and diverging from base
of Rs. Mo, Mg, and Cu, from posterior angle of cell, their basal parts curved and
approximated. Cu, from cell somewhat basad of end. Ist A absent. 2nd A strong. Anal
loop large and complete.

Hind wing with Sc and R, anastomosed for some distance. Sc and M, separate or
short-stalked. Cell less than half length of wing. Discocellular weak, its anterior part
concave, its posterior limb strongly oblique. Mo, M, and Cu, from posterior angle of
cell; M, and M, with basal parts curved and strongly approximated; basal part of Cu,
weakly curved and slightly approximated to that of Mg. Cu, from cell at about two-thirds
from base. Three anals present.

Male genitalia. Uncus subtriangular, bilobed, not or weakly setose. Gnathos with
lateral arms arising from tegumen, median element spike-like, with weak terminal
denticles. Transtilla angled dorsad medially. Juxta large and weakly sclerotized, flanked
by coremata bearing strong, specialized scales. Vinculum ventrally broadly rounded,
medially expanded. Valve oval, with complex basal sclerotization and strong subcostal
ridge; distal part fluted and sometimes with somewhat specialized terminal setae.

Female genitalia. Ovipositor with elongate sparsely setose lobes. Posterior apophysis
with cross-bar set at a shallow angle to shaft but not expanded; shaft long and slender,
but strong. Eighth segment long, slender, cylindrical; tergite more or less rectangular, its
anterior margin slightly excised and strengthened. Anterior apophysis as long as posterior,
with triangular sub-basal flange. Ostium unarmed. Ductus bursae slender, membranous,
with a sclerotized collar near bursa. Bursa globular or oval, membranous, with unevenly
distributed spinules.

Life history. Unknown, but the ovipositor appears adapted to inserting eggs into plant
tissues or crevices. The larva of the closely related Decelia terrosalis Snellen bores in
mango seeds.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974 PLAAT 1

Plate 1. Fig. 1—9. New Guinea Odontiinae. 1, Clupeosoma rufistriatum Hampson, &; 2, Neogenesis flaviplagialis Hampson, 4; 3, Trigonoorda

triangularis spec. nov, holotype, & ; 4, Trigonoorda iebelealis spec. nov, paratype, à ; 5, Pseudonoorda minor spec. nov., holotype, & ; 6, Pseudonoor-

da nigropunctalis (Hampson), 6; 7, Pseudonoorda brunneiflava spec. nov, holotype, &; 8, Hyalinarcha hyalina (Hampson), &; 9, Thesaurica noto-
dontalis (Hampson), 4

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1
ge" —

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974 PLAAT 2

Plate 2.

Fig. 10—12. New Guinea Odontiinae, & genitalia. 10, Clupeosoma rufistriatum Hampson; 11,
Neogenesis flaviplagialis Hampson; 12, Trigonoorda triangularis spec. nov.

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974 PLAAT 3

Plate 3.
Fig. 13—15. New Guinea Odontiinae, 4 genitalia 13, Trigonoorda iebelealis spec, nov.; 14.

Pseudonoorda minor spec. nov.; 15, Pseudonoorda nigropunctalis (Hampson)

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974 PLAAT 4

Plate 4.

Fig. 16—18. New Guinea Odontiinae, 4 genitalia. 16, Pseudonoorda brunneiflava spec. nov.; 17,
Hyalinarcha hyalina (Hampson); 18, Thesaurica notodontalis (Hampson)

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1

PLAAT 5

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974
Si sid
AS bit

N Vi -
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Plate 5.

Fig. 19-22. New Guinea Odontiinae, 9 genitalia. 19, Neogenesis flaviplagialis Hampson; 20,
Trigonoorda triangularis spec. nov.; 21, Trigonoorda iebelealis spec. nov.; 22, Pseudonoorda minor
spec. nov.

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1

dann ns on

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 33

Remarks. This genus contains a number of species which collectively range from Samoa
through tropical Asia to Africa. They have been put in C/upeosoma in the Nymphulinae
when authors noticed the stalking of R, with R34 4 or in Noorda when they did not.
The most closely related named genera are: Decelia Snellen, unplaced by Hampson,
though he put one of the nominal species in Noorda; Autocharis Swinhoe (see Amsel,
1965, 1970), whose species were also included by Hampson in Noorda; and Noordodes
Hampson, placed by him in Endotrichinae. Decelza differs in having the third segment
of the labial palpus clearly marked off from the second, not hidden in its scales, in
having the tip of the forewing curved downward, in having the abdomen produced some
distance beyond the anal angle of the hind wing, in having specialized scale-tufts on the
front and middle legs of the male, in having the ovipositor lobes and eighth tergite of
the females less elongate and in having the bursa copulatrix coiled. Autocharis has vein
Ry of the forewing free, not stalked with R; + 4; the cell of the forewing is long and
the discocellular is only weakly curved; in the hind wing veins M, and M, are stalked.

In addition to the type-species at least the following described species belong to this
genus: Noorda distigmalis Hampson, from Africa, Clupeosoma noordimimalis Hampson,
from Africa, Noorda nigropunctalis Hampson, from Perak; Clupeosoma brunneifusalis
Hampson, from New Guinea, Clupeosoma photina Tams and C. lampra Tams, from
Samoa; and Noorda metalloma Lower, from Australia and New Guinea, if the last species
is correctly identified at the British Museum. All these become new combinations in
Pseudonoorda.

Pseudonoorda minor spec. nov.
Bigs!) > las 22

External characters. Frons fuscous, with a sharply defined, clay-coloured, postero-
median triangle. Vertex ferrugineous. Labial palpus fuscous; base beneath contrastingly
light grey. Eye and ocellus fuscous. Antenna dull buff. Thorax above ferrugineous
anteriorly, shading to clay-colour posteriorly. Abdomen above brownish clay-colour, with
posterior margins of segments light grey. Body beneath and legs grey, coxae, femora and
front tibia infuscated.

Forewing above translucent greyish fuscous to postmedial line, brown-tinged beyond.
Costa narrowly dark fuscous. Weak traces of an outwardly oblique, diffuse, dark-fuscous
antemedial line. A weak fuscous streak on discocellular, strongest at anterior and posterior
angles of cell. Postmedial line fine, fuscous, somewhat irregular, denticulate basad on
veins, the denticulations somewhat accentuated. A diffuse, inconspicuous, dark terminal
line, with faint dark rays extending a short distance basad on and between veins. Fringe
fuscous, with a yellowish-buff line at extreme base, followed by a darker line.

Hind wing with a wedge from M to anal fold coloured like forewing; costal and
anal areas pale translucent yellowish grey. No antemedial or discal markings. Postmedial
line obsolescent anteriorly, generally straight but slightly denticulate from M, to Ist A,
there nearly touching termen and ending abruptly. Fringe as on forewing.

Wings beneath as on upperside but shinier, less strongly marked and with terminal
area hardly differentiated in colour.

Expanse 18 mm.

Male genitalia. Uncus about 21/ times as long as wide. Gnathos with median element
rod-like, somewhat decurved and spinose at tip. Marginal scales of corema flanking
juxta rather slender, in strong bunches. Valve irregular in shape, both dorsal and ventral

34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974

margins somewhat incised before tip; subcostal ridge sinuous, rather weak; fluting strong.
Penis rather thick, with an imbricated group of small spine-like cornuti.

Female genitalia. As described for the genus.

Life history. Unknown.

Types. Holotype, g', and 1 & paratype, Mist Camp, 1600 m, 18 Jan. 1939, L. J.
Toxopeus. Allotype, 9, same locality and collector, 11 Jan. 1939. 1 g' paratype,
Bernhard Camp B, 100 m, 10 April 1939, L. J. Toxopeus. Holotype, allotype and one
paratype in Rijksmuseum van Natuurlijke Historie, Leiden; one paratype in Canadian
National Collection, Type No. 12.863.

Remarks. This species closely resembles P. brunneifusalis (Hampson), from the
Oetakwa River, Snow Mountains, but is appreciably smaller (18 as against 22 mm) and
much paler than the holotype of N. brunneifusalis. I think it better to classify the
present series as a distinct species.

Pseudonoorda nigropunctalis (Hampson) comb. nov.
Figs. 6, 15

Noorda nigropunctalis Hampson, 1899: 222.

The present collection contains two specimens and the Canadian National Collection |
has one from Aiyura, Eastern Highlands, New Guinea, ca. 1850 m, which agree well
with Hampson's types in spite of the different provenance of the latter.

1 2 (without abdomen), Araucaria Camp, 800 m, 21 March 1939, L. J. Toxopeus;
1 g, Iebele Camp, 2250 m, 28 Nov. 1938, L. J. Toxopeus.

Pseudonoorda brunneiflava spec. nov.
Eps. 16

External characters. Frons yellow to yellowish buff, with black anterolateral wedges.
Vertex yellow or yellowish buff. Labial palpus black, white at base beneath. Maxillary
palpus black. Basal scaling of proboscis white. Eye and ocellus fuscous. Antenna buff,
dorsally with dark-brown segmental stripes. A fuscous stripe from eye to wing-base.
Thorax above yellow.

Abdomen above light orange-buff, with posterior margins of segments yellow. Body
beneath and legs whitish buff; front and middle tibiae with weak fuscous bands.

Forewing above yellow. Costa to cell contrastingly fuscous with blue metallic lustre.
Black antemedial dot, orbicular dash and comma-shaped discocellular bar, all adjoining
costal stripe. Postmedial line faint, fuscous, denticulate inward with black accentuation
on veins, oblique distad from costal stripe to cell R;, thence regularly convex to termen
before tornus. Whole area beyond postmedial line evenly copper brown with purple
lustre, except slightly paler just beyond postmedial line anteriorly. A row of fuscous
terminal dots between veins. Fringe shining fuscous with a slightly darker line just basad
of middle.

Hind wing above coloured like forewing, but with costal area whitish and its
maculation obsolescent. No markings basad of postmedial line. The latter almost straight
from M, to Ist A, there ending, but succeeded by a weak dark terminal smudge in anal
area. Fringe as on forewing, but becoming paler toward anal area.

Wings beneath as above but somewhat suffused with brownish and with dark areas
considerably less contrasting.

Expanse 15 to 17 mm.

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 35

Male genitalia. Uncus about twice as long as wide. Gnathos with median element
tapering somewhat irregularly from base to narrowly rounded tip. Scale tufts of corema
flanking juxta narrow. Valve broadly and rather evenly rounded; subcostal ridge straight;
fluting weak. Penis straight and weakly sclerotized.

Female genitalia. Unknown.

Life history. Unknown.

Types. Holotype, &,and 1 g° paratype, Araucaria Camp, 800 m, 17 and 19 March
1939, L. J. Toxopeus. Holotype in Rijksmuseum van Natuurlijke Historie, Leiden;
paratype in Canadian National Collection, type no. 12.864.

Remarks. As will be seen from the figures, the genitalia of this species differ con-
siderably from those of P. #igropunctalis, especially in the shape of the valve. A species
identified as P. metalloma (Lower) in the British Museum (Natural History) and
represented in the Canadian National Collection from Padwe, Sepik District, New
Guinea, is somewhat similar, but has the terminal area purplish fuscous and the post-
medial line of the forewing evenly curved. Another, apparently undescribed, species from
Papua and New Guinea has the postmedial line differently shaped but otherwise is very
close to P. brunneiflava.

Hyalinarcha gen. nov.

Type-species Boeotarcha hyalina Hampson.

External characters. Frons flat and oblique, smoothly scaled, somewhat produced, but
not conical. Vertex short, with rather small erect scales. Behind occiput a collar of large,
wide, smooth scales, probably arising from neck. Labial palpus weakly decumbent, with
broad, compressed, distally pointed scaling, exceeding frons by somewhat more than
length of cranium, the segments with their scaling not visibly divided. Maxillary palpus
prominent, with triangularly dilated distal tuft of scales. Proboscis moderately well
developed, scaled at base. Eye large. Ocellus well developed, separated by a considerable
distance from eye. Chaetosema marked by a tuft of slender scales. Antenna of male
filiform, dorsally scaled, ventrally fasciculate. Body rather short and thick. Legs mod-
erately slender, outer tibial spurs of male about half length of inner. Praecinctorium
with simple, transverse, rounded distal scale tuft.

Male genitalia. Uncus flattened, pyriform, the sides downcurved and more heavily
setose than the median part. Gnathos with wide lateral arms and heavy, somewhat trough-
shaped median piece, the latter shorter than uncus and with rows of spines on dorsal
surface distally. Juxta large, pyriform, dorsally rounded. Vinculum shallow, ventrally
broadly flattened, bearing a moderately developed scale-like structure on each side, flanked
mesally and ectally by narrower striated sclerites. Valve of moderate size, weakly fluted;
a very strong subcostal sclerotized bar, ending in a prominent, free, spine-like process
extending beyond body of valve; dorsad of this a weak zone of setae on distal part of
costa; sacculus broadly sclerotized, bounded dorsally by a weak ridge. Penis thick, curved,
with spirally pleated vesica and with a group of very heavily sclerotized spine-like cornuti,
large, few and in a single row distally, becoming small, numerous and multiserial basally.

Female genitalia. Unknown.

Life history. Unknown.

Remarks. Although at present I propose this genus for the type-species only, there
are additional closely related species, which may have to be placed here when the group

36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974

has been studied more comprehensively. The type-species is recorded only from West
Irian, but its relatives range through the Indo-Papuan region.

Hyalinarcha hyalina (Hampson) comb. nov.
Figs. 8, 17

Boeotarcha hyalina Hampson, 1913: 527.

This species is represented in the present collection by a single male, which exactly
matches the type male from the Mimika River in the British Museum (Natural History).
Though Hampson may have meant to designate a holotype, his citation is ambiguous,
reading “2 g' type”. I therefore designate as lectotype the specimen in the British
Museum (Natural History) with the label “Boeotarcha hyalina type g'. Hmpsn”.

1 6, Hollandia, Brinkmann.

Thesaurica Turner

Thesaurica Turner, 1915: 49.

Type-species Sameodes argentifera Hampson, 1913, from Queensland, by monotypy
and original designation.

This genus was erected for a single Australian species, but Hampson correctly asso-
ciated with this in the British Museum collection the additional species Sameodes noto-
dontalis Hampson, from Sandakan, Borneo, and Noorda accensalis Swinhoe, from Thai-
land. The generic transfers appear not to have been published, and these species ac-
cordingly become Thesaurica notodontalis (Hampson) comb. nov, and Thesaurica
accensalis (Swinhoe) comb. nov. One of these is represented in the present collection.

Thesaurica notodontalis (Hampson)
Figs. 9, 18

Sameodes notodontalis Hampson, 1899: 175.

Epipagis notodontalis: Klima, 1939: 316.

The exact identity of this species is uncertain, as topotypical material is not available
at the time of writing and there are at least three superficially very similar species in
New Guinea. The species represented in the present collection is at least closely similar
to T. notodontalis. There are three specimens in the collection.

2 G', Bernhard Camp, 100 m, 10 and 12 April 1939, L. J. Toxopeus; 1 g Araucaria
Camp, 800 m, 28 March 1939, L. J. Toxopeus.

ACKNOWLEDGEMENTS

My thanks are due to Dr. A. Diakonoff and to the authorities of the Rijksmuseum van
Natuurlijke Historie, Leiden, for their patience in allowing me to retain this material
for study for a very long period of time. It would have been impossible to work out the
taxonomy without the continued support of the British Museum (Natural History) and
especially the active help of Messrs. P. E. S. Whalley and Michael Shaffer. The photo-
graphs were made by Messrs. Orin Hanright and Tom Stovell of the Bio-Graphic Unit,
Research Branch, Canada Department of Agriculture, with the help of Mr. D. H. Kritsch.
Mr. Kritsch also made preparations of genitalia and rendered other technical assistance.

E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 37

LITERATURE

AMSEL, H. G., 1965. — Pyraliden-Studien 2. Die Noorda blitealis Wlk.-Gruppe (Lep.: Pyralidae).

— Stuttgarter Beitr. zur Naturk. 144: 1—7.
, 1970. — Afghanische Pyraliden (Lepidoptera: Pyralidae). — Beitr. natürk. Forsch.

Südwestdeutschl. 29: 25—66.

ARCHBOLD, R., A. L. RAND, & L. J. BRASS, 1942. — Results of the Archbold Expeditions. No. 41,
Summary of the 1938—39 New Guinea Expedition. — Bull. Amer. Mus. nat. Hist.
79: 197—288.

Brass, L. J., 1941. — The 1938—1939 expedition to the Snow Mountains, Netherlands New
Guinea. — J. Arnold Arboretum 22: 271—342.

HAMPSON, Sir G. F., 1897.— On the classification of two subfamilies of moths of the family
Pyralidae, the Hydrocampinae and Scoparianae. — Trans. ent. Soc. London 1897:
127—240.

—, 1899. — A revision of the moths of the subfamiliy Pyraustinae and family Pyralidae.
Part II. — Proc. zool. Soc. London 1899: 172—291.

, 1907. — Descriptions of new Pyralidae of the subfamilies Hydrocampinae and
Scoparianae [concluded}. — Ann. Mag. nat. Hist. (7) 19: 1—24.
, 1913. — Descriptions of new species of Pyralidae of the subfamily Pyraustinae [con-
tinued}. — Ann. Mag. nat. Hist. (8) 11: 322—342.
—————,, 1917. — Descriptions of new Pyralidae of the subfamilies Hydrocampinae, Scoparia-

nae, &c. [concluded}. — Ann. Mag. nat. Hist. (8) 20: 265—282.

Kuma, A., 1939. — Pyralididae: Subfam. Pyraustinae II. — Lepidopterorum Catalogus. Pars 94.

MUNROE, E., 1961. — Synopsis of the North American Odontiinae, with descriptions of new genera
and species (Lepidoptera: Pyralidae). — Can. Ent. Suppl. 24: 1—93.

SNELLEN, P. C. T., 1880. — Nieuwe Pyraliden op het eiland Celebes gevonden door Mr. M. C.
Piepers. — Tijdschr. v. Ent. 23: 198—250.

Toxopeus, L. J. 1940. — Nederlandsch-Indisch-Amerikaansche expeditie naar Nederlandsch Nieuw
Guinea (3e Archbold-Expeditie naar Nieuw Guinea 1938—1939): lijst van verzamel-
stations. — Treubia 17: 271—279.

TURNER, A. J., 1915. — Studies in Australian Lepidoptera. — Proc. roy. Soc. Queensland 27:
11-57.

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| J. P. VAN LrrH. — Revision of the Psenini of the Ethiopian Region, including Malagasy
a (Hymenoptera, Sphecidae, Pemphredoninae), p. 39—101, fig. 1—78.

| | Tijdschrift voor Entomologie, deel 117, afl. 3 Gepubliceerd 23-VIII-1974

oe ee ay

or:

REVISION OF THE PSENINI OF THE ETHIOPIAN REGION,
INCLUDING MALAGASY
(HYMENOPTERA, SPHECIDAE, PEMPHREDONINAE)
by
J. P. VAN LITH

Rotterdam

ABSTRACT

The following new forms are described and illustrated: Psen (Psen) leclercgi (9); Psen
(Mimesa) empeyi (4); Psenulus alveolatus (4), P. aurifasciatus (Q), P. bidentatus pallidus (9
and 8), P. freetownensis (2 and 3), P. jacoti (9 and &), P. leoninus (2), P. oweni (9 and
4), P. rugifrons (2), P. sapobaensis (2), P. uelleburgi (2). A supplementary description of
earlier named species is given and males of some of these species are described for the first time.
The subgeneric position of the species of Psen and the relationships of the species of Psenulus are
discussed. Provisional keys and a list of Ethiopian species are presented.

A few years ago Dr. H. Townes, Ann Arbor, kindly sent me about sixty African
Psenini, mainly from Sierra Leone, for identification. Usually material of this group is
not found in great numbers. This relatively large collection therefore stimulated the
undertaking of a new revision of the Ethiopian Psenini.

In 1961 Prof. J. Leclercq published his catalogue of the species of the Ethiopian
region including Malagasy. In that paper of fundamental importance, he recorded four
species of the subgenus Psen and twenty species of Psenulus, of which ten were newly de-
scribed. A description of Psenulus garambae (Leclercq, 1961b) followed soon after-
wards.

The study of nearly all types or paratypes of known species, along with some small
fresh collections, enabled me to give supplementary descriptions, where necessary.
Furthermore, eleven new species and one subspecies have been added and the systematic
status of a few forms, regarded as subspecies by Leclercq, has been reconsidered. At
present 41 species and subspecies of Psenini are known from the Ethiopian region.

In all about 450 specimens have now been recorded, the majority of which I had at
my disposal. The material not seen by me includes a few large series of the commoner
species, like a series of over sixty Psenulus capensis from Garamba. It is evident that the
above total is far from being representative for this large area. Nevertheless we have
obtained a somewhat better idea of its Psenine fauna. Some species have dispersed over
large areas, consisting of savanna as well as forest regions. Psenulus capensis Brauns, for
instance, is found from the extreme South of Africa to as far north as Sierra Leone and
Ethiopia, with very little variation in structure or colour. Of Psenulus bidentatus (Came-
ron) at least three subspecies can be distinguished. The nominate form occurs mainly in
the savanna region, pallidus subsp. nov. in the forest region and the subspecies rabrocau-
datus Turner in South Africa.

The subgenus Mimesa was already known from North Africa; the first representative
of this subgenus of Psen south of the Sahara has now been found in Transvaal. The

40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

subgeneric status of a few species, thus far thought to belong to Psen s.str., is discussed
at the end of the present paper, where also an attempt is made to recognize some higher
groups of Psenulus.

The few data we have concerning the bionomics of the Ethiopian Psenini, their prey
(cf. Psen silvaticus Arnold, Psen madecassus Arnold, Psenulus stevensoni Arnold and
Psenulus capensis Brauns) or their nesting (cf. Psenulus patei Arnold and Psenulus
capensis) agree with our present knowledge of the bionomics of their Palaearctic or Indo-
Australian relatives.

For generously sending me their material I am much indebted to the authorities and
staff of the institutions as well as to the private entomologists mentioned below, preceded
by abbreviations used in the following text:

AMG — Albany Museum, Grahamstown, Cape Province, South Africa; Dr. F. W.
Gess

AMNH — The American Museum of Natural History, New York, U.S.A.; Dr. J. G.
Rozen and Mrs. M. Favreau

BM — British Museum (Natural History), London, United Kingdom; Dr. I. H. H.
Yarrow and Mr. C. R. Vardy

CNC — Canadian National Collection of Insects, Ottawa, Canada; Dr. C. M. Yoshi-
moto

EC — Collection H. N. Empey, Highlands North, Transvaal, South Africa

FAG — Faculté des Sciences Agronomiques de l'Etat, Gembloux, Belgium; Prof. J.
Leclercq

HT — Collection H. Townes, Ann Arbor, U.S.A.

IRSNB — Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium; Dr. P.
Dessart

MRAC — Musée Royal de l’Afrique Centrale, Tervuren, Belgium; Dr. J. Decelle

MS — Naturhistoriska Riksmuseet, Stockholm, Sweden; Dr. P. I. Persson

NMB — Naturhistorisches Museum, Basle, Switzerland; Dr. C. Baroni Urbani

OSUM — Oregon State University, Entomological Museum, Corvallis, U.S.A.; Dr.

G. R. Ferguson
PMFV — Collection P. M. F. Verhoeff, Utrecht, The Netherlands

RM — National Museum of Rhodesia, Bulawayo, Rhodesia; Mr. F. C. de Moor

SAM — South African Museum, Cape Town, South Africa; Dr. A. J. Hesse

TM — The Transvaal Museum, Pretoria, Transvaal, South Africa; Dr. J. A. van
Reenen

USNM — National Museum of Natural History, Smithsonian Institution, Washington
D.C., U.S.A; Dr. K. V. Krombein, Dr. P. D. Hurd and Dr. A. S. Menke

ZMB — Zoologisches Museum der Humboldt-Universität, Berlin, D.D.R.; Dr. E.
Königsmann

Special thanks are due to Prof. Jean Leclercq, Gembloux, and to Mr. F. C. de Moor,
Bulawayo for their assistance. Mr. de Moor kindly also sent me some interesting un-
published notes by Arnold. I am also much obliged to Dr. J. G. Betrem, Deventer, Drs.
J. P. Duffels of the Institute of Taxonomic Zoology, Amsterdam, Mr. H. N. Empey,
Transvaal, Dr. Per Inge Persson of the Natural History Museum, Stockholm, and to the
South African Embassy, The Hague, for courteous aid in securing bibliographic, topo-
graphic or other valuable information.

J. P. vAN LITH: Revision of Ethiopian Psenint 41

KEY TO THE SPECIES OF Psen S.L.

(the females of P. empeyi, P. patellatus, and the males of P. lecleregi and P. silvaticus

are still unknown)

Belles lode Ug PaO TEP Mn 2
MAES iur: TONE POE Pee)
Mesopleura Reledingi pe il area at striato- nance) San densely,
coarsely punctate, interstices narrower than punctures. Petiole dorsally smooth . 3
Mesopleura much finer and more sparsely punctate. Scutum coarsely punctate . 4

Propodeum finely reticulato-carinate, enclosed area ill-defined, more coarsely reticu-
lato-carinate. Frons medially rugoso-punctate. Thorax black, pronotal tubercles and
parts of pronotum red. Length 12 mm. Male unknown. Rhodesia :

Psen (Psen) silvaticus not
peten sa ek carinate, ai area well-defined. Frons densely
coarsely punctate, partly striato-punctate. Thorax largely black, pronotum, pronotal
tubercles, anterior corners and narrow lateral margin of scutum, and upper two-
thirds of anterior en of hora reddish. Length 11.5 mm. Male unknown.
Malagasy . . . mu... Psen (Psen) leclercgi sp. nov.
Frons coarsely striato- tate meg more distinctly and more densely
punctate, interstices 3 or 4 times diameter of puncture. Petiole dorsally smooth.
Pygidial area finely coriaceous with a few irregular punctures along sides. Thorax
and gaster completely black, legs dark. Length 14 mm. Zaire . 5

si Psen (Psen) ao asta
Eh ali stel ate deb finely and very sparsely, indistinctly,
punctate. Scutum coarsely punctate, interstices usually at most as large as punctures.
Petiole with fine latero-dorsal groove, in which a row of punctures, each with erect
long hair. Pygidial area shining, with a few punctures along sides. Thorax including
mesopleura largely dark reddish, gaster including petiole reddish-brown, tibiae
reddish. Length 10— 11 mm. Malagasy . . Psen (subgenus?) madecassus Arnold
Apical margins of third and fourth gastral sternites with fasciculate hairs. Scutum
striato-punctate. Mesopleura distinctly punctate. Black, fore and mid tibiae brownish.
Length 12—13 mm. Zaire . . . . + + . Psen (Psen) congolus Leclercq
Apical margins of third and fone sternites without fasciculate hairs. Thorax
largely dark reddish, or gaster partly reddish, or tarsi whitish. . . . . . . 6
Frontal carina distinct. Mesopleura including hypo-epimeral area finely, very
sparsely, punctate. Petiole with fine latero-dorsal groove, in which row of punctures,
each with erect long hair. Thorax including mesopleura largely dark reddish, tibiae
reddish, i including petiole reddish-brown. Length 8.5—9.5 mm. Malagasy .
3 Psen (subgenus?) madecassus Arnold
Frontal carina ln Hate densely punctate. Thorax black . . . . 7
Hypo-epimeral area dull, coarsely coriaceous. Mesopleura densely punctate, with
narrow interstices. Scutum shining, densely punctate, interstices mostly larger than
punctures. Antennae normal, gradually broadening towards apex. Legs normal.
Ventral plate of petiole, first gastral tergite and base of second tergite reddish.
Length 6.5 mm. Female unknown. South Africa . Psen (Mimesa) empeyi sp. nov.

— Hypo-epimeral area shining, with distinct, separate, punctures. Mesopleura densely

—

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

finely punctate, narrow interstices. Scutum shining, densely somewhat irregularly
punctate, interstices sometimes larger than punctures. Apical half of flagellum
broadened, segments much concave below. Fore femora, fore tibiae and especially
fore basitarsi broadened. Gaster black, all basitarsi whitish. Length 7 mm. Female
unknown. South Africa . . . . . . . Psen (subgenus?) patellatus Arnold

KEY TO THE FEMALES OF Psenulus
(the females of P. alveolatus, P. avernus and P. trevirus are still unknown)

Tempora with coarse sculpture. Interantennal carina much broadened. Transverse
carina below antennae. Prescutal sutures long, on apical half at least slightly indi-
cated. Propodeal enclosure with wide central area. Petiole usually short, cylindrical,
base flattened. No distinct pygidial area. Apical margins of sternites 4—5 with
dense fringe of long pale hairs . . . . . . (Group of Psenulus turneri) 2
Tempora with much finer sculpture, in some species finely striate. Sternites 4—5 on
apical margin with short pubescence, or with fringes of medium length (see

leoninus; veticulosas and sapobaensas). culi Pils © NE, ST, ADITO
Second submarginal cell of fore wings petiolate. . . . u
Second submarginal cell of fore wings not petiolate, if alan see aso turnert or
paulisae . . 5 5

Apical part of fourth belgie Blas: ee zes NA (ad also So ui
tergite. Fore legs reddish, mid legs more or less brown, hind legs black. Antennae
dorsally reddish-brown. Frons obliquely striato-punctate, vertex punctate with faint
striae. Scutum rather coarsely punctate with traces of transverse striae. Propodeum
behind enclosure irregularly striate. Mesopleura strongly punctate with tendency to

striation. South Africa. . . . . twrneri Arnold
Tergites 1—6 or 2—6 reddish. an Blackich don all At least base of hind
tibiae reddish. Upper part of propodeum obliquely striate . . . . . . . 4

Frons below ocelli distinctly obliquely striato-punctate. Vertex shining, finely
punctate. Clypeus distinctly bidentate. Mesopleura and scutum distinctly densely
punctate with tendency to transverse striation. Fore femora yellowish-red, sometimes
with brown streak behind; mid femora brown behind; hind femora black, hind
tibiae dorsally brown with pale reddish base. Zaire, Sierra Leone . pazlisae Leclercq
Frons finely densely punctate, slightly striate near median carina. Vertex shining,
very finely punctate. Clypeal margin slightly emarginate, teeth less distinct. Scutum
and mesopleura much finer and more sparsely punctate. Fore and mid legs and also
hind tibiae yellowish-red; hind femora partly brown. Male unknown. Zaire, Sierra
L'CONcH ARA «ll. | perusasiLecleteq
Second racial cl i o wines Ario or Ron so (see also twrneri and
paulisae); if slightly open, at least mid basitarsi pale yellowish. Upper part of
propodeum striate. Frons more or less obliquely striato-punctate, or distinctly
punctate. Tergites 1—2 or 1—3 black. . . . . step En
Second submarginal cell distinctly open dorsally (see also: Par me saltitans). Frons
striato-punctate or finely punctate. Gaster completely black or tergites 1—6 red, or
upper part of propodeum coarsely reticulate. . . . aak 8
Apical margin of second tergite and tergites 3—6 completely ea Legs cu
greater part of all femora dark brown, also back of mid tibiae and hind tibiae; mid

10.

bts

J. P. vAN LITH: Revision of Ethiopian Psenini 43

basitarsi yellowish-brown; hind tibiae with yellowish basal ring. Antennae slightly
darkened dorsally. Mandibles yellowish-red. Vertex shining. Scutum shining, finely
punctate, anteriorly somewhat striato-punctate. Mesopleura distinctly punctate, more
sparsely than in paulisae, interstices smaller than punctures. Male unknown. Zaire

garambae Leclercq
Tele Dig radish den most allo ek oF Ti ito) rar) bte Add ER

Second submarginal cell usually triangular or nearly so. Scutum shining, finely
punctate, laterally with tendency to transverse striation. Tergites 4—6 reddish. Fore
legs yellowish-red, underside of femora and trochanters brown; mid femora dark
brown, underside of mid tibiae pale reddish-brown, basal fifth yellowish; hind
femora and tibiae brownish-black, basal fourth of hind tibiae straw-yellow; all
basitarsi pale yellowish, apex of hind basitarsi more or less darkened. Antennae
dorsally slightly brownish. Male unknown. Rhodesia. . . . . . pate? Arnold
Second submarginal cell not completely closed. Scutum shining, slightly stronger, but
still finely punctate, with very little tendency to transverse striation. Apical margin
of third tergite and whole tergites 4—6 reddish. Fore femora almost completely
brown, fore tibiae yellowish-red, fore tarsi yellowish; mid femora and tibiae brown,
mid basitarsus pale yellowish; hind legs brown, tibiae without distinct basal ring.
Antennae brownish dorsally. Male unknown. Rhodesia . . . . saltitans Arnold

Gaster black, last tergite brownish. Legs black, fore tibiae, apex of fore femora and
back of mid tibiae reddish-brown. Frons on either side of median carina convex,
shining, very finely and sparsely punctate. Scutum shining, finely sparsely punctate.
Prescutal sutures indistinct on posterior three-quarters part of scutum, indicated only
by a line of punctures. Mesopleura distinctly punctate, interstices mostly larger than
punctures. Upper part of back of propodeum finely obliquely striate. Larger species,
length 8 mm. Petiole about one and one-quarter times as long as first tergite, in
dorsal aspect. Male unknown. Sierra Leone . . . . . . aurifasciatus sp. nov.
At least tergites 5—6 reddish . . . . a Bte
Tergites 1—6 reddish. All legs including pira tok i, ini Antennae
dark brown above; scape, last segment completely and rest of segments below
yellowish-brown. Frons, scutum and mesopleura finely punctate. Upper part of
propodeum smooth and shining. Petiole shorter than first tergite, in dorsal aspect.
Sierra Leone, Equatorial Guinea . . . . freetownensis sp. nov.
Only tergites 4—6 or 5—6 completely il ie lest mid and hind femora brown,
fore tibiae and foreside of mid tibiae reddish, hind tibiae brown with yellowish-grey
base. Antennae dorsally blackish-brown, scape and antennae below brownish. Frons
above antennal sockets obliquely striato-rugose. Scutum and mesopleura striato-
punctate. Upper part of propodeum coarsely reticulate, laterally somewhat obliquely

striate. Petiole about as long as first tergite. Zaire, Sierra Leone . . ealae Leclercq
First gastral tergite not distinctly nodose; or thorax with Ka 5 marking and
gaster reddish or partly reddish-brown . . Le Ze

First gastral tergite nodose. Frontal carina narrow bebe antennae. No sharp
transverse carina below antennae, at most a low ridge. Anterior margin of clypeus
blunt. Prescutal sutures short. Thorax and Di black. (Geo of Psenulus latian-
mula) (ats: 4 30
Raised part of nanna carina more or re Binden sometimes not excavate.

12)

14.

19%

16.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

Prescutal sutures long (except in reticulosus), sometimes partly indistinct. Some
species with reddish petiole and gaster . . Ana TZ
Raised part of interantennal carina narrow Ge slightly ied in dilectus)
between antennae. Prescutal sutures short. Never gaster including petiole reddish

Gaster and petiole reddish. Fore and mid legs whitish or yellowish. Distinct trans-
verse carina below antennae . . . Larga DE)
At least petiole black, if RES andy fie cate gite Ch Bess, legs not
whitish or yellowish . . . . “nes o)

e part of carina between antennae eh aan ratio Sort length about

: 1.5. Prescutal sutures long but not sharply defined. Scutum shining, distinct,
Ro punctation, sometimes with tendency to rugosity. Upper part of back of
propodeum smooth. Petiole about one and one-third times as long as first tergite, in
dorsal aspect. Pygidial area distinct, narrow. Pronotal dorsum, pronotal tubercles,
hind margin of scutellum and metanotum ivory-white. Fore and mid legs including
coxae ivory-white; hind = yellowish-red, hind trochanters more or less yellowish-

white. Malagasy . . . + . fulgidus Arnold
Carina between antennae een iii sometimes Maltees: sharp. Petiole over
one and a half times as long as first tergite, in dorsal aspect. . . . . . . 14

Prescutal sutures distinct only on anterior third of scutum. Scutum with distinct
separate punctures. Propodeum behind enclosure smooth, back reticulate, above with
oblique striae. Pygidial area narrow, with parallel unsharp lateral carinae. Pronotum
yellow dorsally, in var. alboscutellatus Arnold also scutellum and metanotum yellow.
Fore and mid legs including trochanters yellowish, hind legs reddish. Antennae
reddish, base dorsally slightly darkened. Malagasy. . . . . reticulosus Arnold
Prescutal sutures distinct on about two-fifths part of scutum, continued by a row of
punctures. Scutum with variable, distinct separate punctures. Behind propodeal
enclosure narrow smooth margin, back-side obliquely striate. No pygidial area. Mid
tibiae on outer apex with longitudinal groove. Pronotum medially and metanotum
somewhat brownish (discoloured?). Fore and mid legs including trochanters and
mid coxae straw-yellow, hind legs including coxae reddish. Antennae dorsally dark
brown, last segment above and flagellum below reddish-brown. Male unknown.
INSERT IA NE STE sapobaensis sp. nov.
Apex of iu Sini Sigi ans and Tone a longitudinal carina. Frontal
carina between antennae only slightly broadened. Frons finely punctate. Distinct
transverse carina below antennae. Prescutal sutures long. Back of propodeum smooth.
Petiole about one and a half times as long as first tergite, in dorsal aspect. Petiole
except ventral plate black, gaster reddish. Pronotal tubercles and tegulae yellowish-
red. Fore and mid legs except back of femora and base of hind tibiae yellowish,
hind femora and apex of hind tibiae brown. Zaire . . . . . benoiti Leclercq
Pygidium without single longitudinal carina. . . NE EURO
No transverse carina below antennae. Broadened part Fd raced tarten carina
indistinctly closed below. Protruding median part of clypeal margin slightly emargi-
nate, almost blunt. Frons and vertex densely finely punctate. Scutum densely finely
punctate, slightly transversely rugose, prescutal sutures with short transverse striae
and deep alveoles. Petiole at least one and a half times as long as first gastral tergite,
in dorsal aspect. Pygidial area elongate-triangular. Gaster black. All tibiae and tarsi

197%

18.

19.

20.

21e

20%

J. P. van LITH: Revision of Ethiopian Psenini 45

whitish; fore and mid tibiae below and apical half of tibiae, except a narrow line

above, brownish. Male unknown. Rhodesia. . . . ale. bob Arnold
Distinct transverse carina below antennae, sometimes fine and only visible in ventral
aspect. At least tergites 5—6 reddish or hind tibiae dark. . . . 17

Petiole little longer than first tergite. Apical margins of fourth aad) fifth one
with fringe of yellowish-golden hairs, at least as long as width of base of hind
basitarsi, but shorter than in group of turneri. Frontal carina ending normally as
single carina in transverse carina below antennae, transverse carina almost rectangular
in frontal aspect. Scutum partly transversely striato-punctate and also with some very
large punctures. Prescutal sutures ill-defined. Apical half of pygidial area narrow
with parallel carinae. Apical margin of first tergite and tergites 2—6 completely
reddish. Legs reddish, back of mid femora and hind femora except underside dark
brown. Fore and mid basitarsi GET Length about 6 mm. Male unknown. Sierra

Benet OU. Sule hee) eon nis sp. NOV:
Petiole much lonen Apical margins 1. fourth i fifth sternites without fringes of
long hairs. . . ls

Raised part of Bone) carina ) meh Broadenedt betweet aften proadened part
usually not closed below but ending with two parallel carinae in transverse carina
below antennae. Upper part of propodeum finely obliquely striate. Pygidial area
distinct. Fore and mid legs not whitish, at most mid basitarsi . . 5 dl)
Carina between antennae much less broadened, or broadened part drat and inter-
antennal carina ending as a single carina in transverse carina. Lateral carinae of
pygidial area not well-defined, if more distinct, fore and mid legs extensively

whitish. Petiole about twice as long as first tergite, in dorsal aspect . . . . 23
Gaster black, at most last tergite brownish, or petiole completely and first tergite
parilygredcish-browntt Mt... 2,2,.2,2 2 ae es dien 20
Gaster partly reddish . . . . #21

Frons very finely obliquely striate, vertex ih call sculptures Scum del super-
ficially transversely striate and with some larger punctures. Petiole more than twice
as long as first tergite, in dorsal aspect. Antennae reddish, apical half darkened. Fore
and mid legs reddish-brown; at least mid basitarsi yellowish-white; hind legs entirely
brown. Pronotal tubercles blackish; black. Length 7 mm. Rhodesia .

3 stevensoni Arnold
Ea ona finely ee, ek mat meer Scutum shining, anterior
two-thirds sparsely punctate. Petiole less than twice as long as first tergite. Antennae
reddish-brown, segments 7—11 more or less darkened dorsally. Fore and mid legs
including mid basitarsi and base of hind tibiae reddish-brown; fore and mid femora
partly darkened, hind femora black, hind tibiae dark brown. Pronotal tubercles
reddish-brown. Gaster black, petiole at least ventrally and laterally partly reddish-
brown, also first tergite more or less brown, apex of last segment yellowish-red.
Length 7 mm. South Africa . . . . . jfacoti sp. nov.
Apical margin of fourth tergite and Late 56 zoeen reddish. Mid basitarsi
brown. Scutum dull, finely punctate. Petiole over twice as long as first tergite, in
dorsal view. Pygidial area broader, lateral carinae slightly converging towards apex.
Length 8 mm. Male unknown. Rhodesia . . . . . . . . luciuosus Arnold
Gastral tergites 1—6 reddish. Pygidial area narrow . . bide 22
Frons very finely punctate, vertex almost impunctate. Petnle over twice as long as

208

24.

25%

26.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

first tergite, in dorsal aspect. Antennal segment 1—12 reddish below. Greater part of
mandibles reddish-brown. Legs brown, fore tibiae and tarsi reddish, mid basitarsi
testaceous. Length 7.75 mm. Male unknown. Zaire. . . . . lusingae Leclercq
Frons finely densely striato-punctate, vertex indistinctly striate. Petiole slightly over
one and three-quarters times as long as first tergite, in dorsal aspect. Antennae black,
only small central part of segments 5—12 below reddish-brown. Mandibles darker.
Legs black, last tarsal segments, underside of fore tibiae and base of hind tibiae

brownish. Length 7 mm. Male unknown. Ethiopia. . . . . rugifrons sp. nov.
Tergites 1—6 or 2—6 reddish . . . Mrt 24
At least tergites 1—3 black or very ane prow mare rad end aaf edi
brown". at MERG

Pronotal ande at most hy reddish or Rare beo ae rarely yellowish;
if so cf. bidentatus pallidus). Fore and mid legs reddish, underside femora brownish,
base of fore tibiae often somewhat yellowish, rarely a narrow yellowish streak on
foreside, fore and mid basitarsi yellowish or yellowish-red; hind legs reddish on
inner side, dark brown on outer side, base of hind tibiae usually dark. Mandibles
reddish. Antennae reddish, apical half dorsally brown. First gastral tergite mostly
dark brown (reddish in female from Angola). Punctation of scutum superficial,
interstices larger than punctures, slight tendency to transverse striation. Propodeum
with narrow smooth margin behind enclosed area, back dorsally obliquely striate,
below irregularly carinate. Pygidial area dull, lateral carinae not sharp, parallel.
Length 8.5—9.5 mm. Tanzania, Kenya, Rhodesia, Angola, Zaire .
ike . bidentatus bidentatus (Cn
At ies: posero: half of TA tubereles Aaa foreside of fore tibiae distinctly
yellowish. Tergites 1—6 or apex of first tergite and tergites 2—6 completely
reddish. Length 7.5—8 mm . . . LIFE RISORSE 25
Only pronotal tubercles, foreside of fare “ao Ba DE aid apie and fore and mid
basitarsi yellowish or whitish. Base of hind tibiae paler than rest of hind tibiae.
Punctation of scutum coarser than in bidentatus bidentatus, more striato-punctate
with interstices larger than punctures and more strongly transversely striate. Sculpture
on back of propodeum reaching enclosed area, except laterally. Pygidial area as in
nominate subspecies. Length 7.5 mm. Zaire, Rhodesia, Malawi, Equatorial Guinea
bidentatus pallidus subsp. nov.
PORC partly, tonen bereide and metanotum yellow. Fore and mid legs
yellowish-white, femora with light brown streak behind; greater part of hind legs
brown. Scutum finely punctate, slight tendency to transverse striation. Propodeum
behind enclosed area smooth, back with oblique striae above and irregular carination
below. Pygidial area dull, more distinctly defined, almost rectangular and with two
rows of fine punctures. ae 8 mm. Male unknown. Nigeria, Sierra Leone .
nigeriae Leclercq
At east cite 56 as Sui as apical half or four rei reddish. Thorax black,
pronotal tubercles partly dark brown. Fore legs except trochanters reddish-brown,
underside of fore femora darker; mid legs reddish-brown, underside of femora and
tibiae dark brown; hind legs dark brown or black. Mandibles reddish. Scutum with
large, somewhat superficial punctures and distinct transverse rugae especially on
outer side of prescutal sutures. Narrow smooth margin behind enclosed area of pro-
podeum, back with fine oblique striae, medially sometimes reaching enclosed area.

27:

28:

29.

30.

J. P. van LITH: Revision of Ethiopian Psenini 47

Pygidial area as in bidentatus bidentatus. Length 8.75 mm. Male unknown. South
ARIANE „IR. . . + . bidentatus rubrocaudatus Turner
Tergites 4—6 geek be Thorax black including pronotal tubercles. Legs some-
what darker reddish-brown, except fore basitarsi. Mandibles reddish. Scutum finer
punctate, very little tendency to transverse striation. Narrow smooth margin behind
enclosed area, back side dorsally with distinct fine oblique striae. Length 9.5 mm.
Male’unknown. Ethiopia . . „Ti et. . . . bidentaius subsp. nov.?

No well-defined pygidial area. Scutum transversely rugoso-punctate. Petiole less
than twice as long as first tergite. This tergite of usual shape. Upper part of pro-
podeum laterally with oblique striae, on either side of median sulcus a smooth area.
Back of propodeum with coarse reticulate carination. Distinct transverse carina
below antennae. Median part of pronotum, and whole metanotum, yellowish. Petiole
black, tergites 1—6 reddish. Fore and mid legs yellow, fore and mid femora brown
below, hind femora completely brown. Antennae dorsally dark brown. Length 7.5

mm. Male unknown. Zaire, Uganda. . . . een wghesgutenenlteclereg
Distinct narrow pygidial area. Scutum oi finely He First gastral tergite
Sliphitly}modoseltt. tu. |. 128

Hind legs and tergites 1—6 een Pronotum JU oi ess small
marks on scutum, scutellum, metanotum, large part of back of propodeum and a
mark on anterior plate of mesepisternum pale yellow, also fore and mid legs. Petiole
black with yellowish base. Transverse carina below antennae lacking or indistinct.
Back of propodeum almost sega smooth. Length 8—9 mm. Sierra Leone

3 oweni sp. nov.
Hind LA ac aie dea ria or ati Sa coe dark brown and only
Apicdemarcinsidancreddishers wah. banusnanl dert de store wer stun mul 29
Pronotum, pronotal tubercles, four marks on scutum, scutellum, metanotum, back of
propodeum and parts of sides of thorax yellow. Base of petiole yellow, remainder
dark brown, gaster brownish. Fore legs mostly yellow; foreside mid legs yellowish,
back-side brownish-red; hind legs brown but back of femora partly yellow. Fine
transverse carina below antennae. Upper part of propodeum smooth, lower two-
thirds of back-side with fine reticulate carination. Petiole about twice as long as first

tergite. Length about 10.5 mm. Male unknown. Equatorial Guinea . 2
uelleburgi si nov.

Pronotum dorsally, pronotal tubercles, scutellum and metanotum yellowish-red.
Petiole reddish-brown or darker brown, base of tergites dark brown, apical margins
dark reddish. Fore and mid legs including trochanters reddish, mid femora brownish
behind, hind legs brown, base of tibiae reddish. Transverse carina below antennae
fine or indistinct, vertical carinae downwards from outer side of antennal sockets
distinct. Propodeum behind enclosed area smooth, back-side finely punctate, apex
finely reticulate. Petiole slightly longer than first tergite. Length 8 mm. Male
unknowg.Malapasy; Eee. honest werd deg ver wll ‘nd air dlectzs Saussure

Lateral corners of pronotum obtuse. No sharp transverse carina below antennae, at
most an indistinct ridge, but below outer side of each antennal socket a short
vertical carina, in dorsal aspect protruding like a tooth. Punctation of scutum
variable, sometimes very coarse. Upper part of back of propodeum usually distinctly
obliquely striate. Pygidial area indistinct. Foreside of fore and mid tibiae yellow, also

Dlt

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

fore and mid basitarsi and base of hind tibiae. Length about 7 mm. South Africa,
Mozambique, Malawi, Zambia, Tanzania, Uganda, Zaire, Sierra Leone. The sub-
species /aevior Arnold, Ethiopia, female unknown, is distinguished by the brown
femora and weaker sculpture of scutum and propodeum . capensis capensis Brauns
Lateral corners of pronotum sharp. No distinct transverse or vertical carinae below
antennae, sometimes a small low shining tubercle below antennae. Punctation of
scutum variable but usually finer than in capensis. Propodeum behind enclosed area
usually smooth. At least foreside of fore and mid tibiae more or less yellowish.
hanger: waal fee de elec We athe Side" ee Se chi a RE
Fore tarsi, a part of mid basitarsi, and base of hind tibiae yellowish-white. Puncta-
tion of scutum variable, usually fine and sparse. Pygidium with two indistinct, close,
parallel carinae at apex. Length 8—9 mm. Tanzania, Rhodesia, Uganda, Zaire,
Nigeria, Sierra Leone. . |. . +. . datiannulatus latiannulatus (Cameron)
Mid basitarsi and base of kia tibiae black or dark brown. Punctation of scutum
stronger. Distinct narrow pygidial area, with very close, parallel carinae. Length 7.75
mm. Zaire, Rwanda, Cameroon. . . . . . latiannulatus basilewskyi Leclercq

KEY TO THE MALES OF Psenulus

(the males of P. aurifasciatus, P. dilectus, P. erusus, P. garambae, P. ghesquierei, P. kobli,
P. leoninus, P. luctuosus, P. lusingae, P. nigeriae, P. patei, P. rugifrons, P. saltitans,

P. sapobaensis, and P. velleburgt are still unknown)

Tempora coarsely, mostly longitudinally, striato-reticulate. Raised part of carina
between antennae more or less broadened. Antennae with tyloidea. Anterior margin
of pronotum alveolate. Prescutal sutures long, almost reaching hind margin of
scutum. At least thorax and petiole black . . . (Group of Psenulus turneri) 2
Tempora much less coarsely sculptured, sometimes finely striate (see benoiti) . 6

Upper part of back of propodeum smooth. Frons shining, finely punctate. Antennal
segments long, segments 3 (or 4)—8 with distinct tyloidea. Scutum shining, finely
punctate. Second submarginal cell of fore wings open dorsally. Tergites 1—6
reddish, only last sternites slightly brownish, all legs reddish. Sierra Leone, Equa-

torial Guinea . . . . freetownensis sp. nov.
Upper part of back a de sail si At least first tergite com-
pletely.blackind. hab sum: pane ahi bone NES

Second submarginal cell not petiolate or triangular, dorsally open. Frons obliquely
striate. Antennal segments 8—12 at least one and a half times as long as broad at
apex, third segment with very small tyloidea, segments 4—10 with distinct long
tyloidea, on third and 11th segment at most a small point. Scutum strongly trans-
versely striate. Petiole about one and a half times as long as first tergite, in dorsal
aspect. Tergites 1—3 black or dark brown. Fore and mid tibiae and tarsi and fore-
side of fore and mid femora reddish; back of fore and mid femora, as well as hind
legs completely, brown except pale yellowish base of hind tibiae. Zaire, Sierra Leone

: ealae Leclercq
Soi Sebra cell idee te ae la in turner? or paulisae). Base of
hind:tibiaeidarkn: Dei yao WIR en SD ai Arr

J. P. van LITH: Revision of Ethiopian Psenint 49

Petiole nearly one and one-third times as long as first tergite, in dorsal aspect. Frons
obliquely striate. Antennae moniliform, segments 8—12 little longer than broad,
segments 3—10 with distinct tyloidea, small on tenth segment. Scutum distinctly
punctate, slight tendency to transverse striation. Prescutal sutures with large alveoles.
First tergite, basal two-thirds of second tergite and sides of tergites 3—5 partly
black. Fore and mid legs including trochanters reddish, also hind tibiae and tarsi,
hind femora brown. Zaire, Sierra Leone m based on male from Sierra

BEONE) = . i 02) 2) paulisaeleelereq
Petiole usually diaz lugo ou one and a ale x or twice as long as first
tergite, in dorsal aspect, or scutum more coarsely sculptured. . . . . . . 5

Petiole about one and a half times as long as first tergite, in dorsal aspect. At least
tergites 1—3 fully black, fourth tergite more or less darkened. Fore legs including
trochanters reddish-brown, femora brown below, mid legs reddish-brown, darkened
below, trochanters brown; hind legs for greater part dark brown. Antennal segments
3—8 about one and a half times as long as wide at apex, segments 4—8 with
distinct small tyloidea, 9—10 with indistinct tyloidea. Frons obliquely striate, vertex
more superficially striato-punctate. Scutum more coarsely punctate than in pazlisae,
with some transverse rugae, especially across the prescutal sutures, and large alveoles.
South Africa . . . woes urnen Arnold
Petiole nearly twice as iene as ae ent Set O and two-thirds of second
tergite black, greater part of sternites 2—4 blackish-brown. Fore legs including
trochanters reddish-brown, mid legs slightly darker brown, hind legs dark brown.
Antennal segments 3—8 about one and a half times as long as wide at apex,
segment 3 with small tyloidea, segments 4—8 with longer, distinct, tyloidea. Frons
and interocellar area coarsely, frons obliquely, striato-punctate, vertex between ocelli
and oculi more finely striato-punctate. Anterior half of pronotal dorsum with
stronger longitudinal short cärinae, alveolate. Scutum distinctly punctate, tendency
to transverse striation, rugae across prescutal sutures very coarse, alveoles larger than
in turneri. Female unknown. Rhodesia . . . Uni... alveolatas'sp. nov.
First gastral tergite not distinctly nodose, or arie 1—6 or 2—6 reddish . . 7
First gastral tergite distinctly nodose. Prescutal sutures short. Raised part of frontal
carina between antennae sharp. Gaster black, fore and mid tibiae at least partly
yellowish or yellowish-white . . . . . (Group of Psenulus latiannulatus) 16
Petiole and gaster reddish. Interantennal carina not distinctly broadened between
antennae, as it is in female. Upper part of back of propodeum smooth behind
enclosed area. Petiole about one and a half times as long as first tergite. . . 8
Petiole black, at most base yellowish, or sides dark reddish-brown. . . . . 9
Scutum coarsely transversely striato-punctate. Mesopleura coarsely punctate, hypo-
epimeral area more finely. No tyloidea on antennal segments. Scape of antennae
yellow, rest of antennae yellowish-brown, brownish above. Pronotum, pronotal tuber-
cles, tegulae, hind margin of scutellum and metanotum yellow. Fore and mid legs
including trochanters yellow; hind legs i coxae a ner Length 6 mm.
Malagasy . . . 5 . . + fulgidus Arnold
Scutum shining, Ca Mare A variable size, interstices mostly larger than
punctures. Mesopleura finely punctate. Antennal segments 3 (or 4)—13 with
distinct oblique tyloidea. Scape of antennae yellow, rest of antennae reddish. Two
marks on pronotum, and pronotal tubercles, yellow or yellowish-white. In var.

10.

12%

13:

14.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

alboscutellatus Arnold also scutellum with yellowish-white mark. Fore and mid legs
including trochanters whitish, femora brownish below and last tarsal segments
reddish; hind legs ee: coxae reddish. Length 8.5 mm. Malagasy .

reticulosus Arnold
mora: ata aast ple at most Sa ped nad sides of petiole reddish-
brown. Interantennal carina broadened. Median part of propodeal enclosure rather
broad. Upper part of back of propodeum finely Eh striate. Petiole long . 10
Gaster more or less reddish . . . CANARIA
Petiole at least twice as long as dt i Hat in dl ar Frons with
superficial oblique striae. Antennal segments 10—12 about as long as broad; seg-
ments 3—13 with tyloidea. Scutum transversely striato-punctate, prescutal sutures
almost reaching hind margin. Pronotal tubercles dark brown. Legs and veins of fore
wings reddish-brown. Petiole completely black. Gaster with somewhat appressed
silvery pubescence. Length 6 mm. Rhodesia. . . . + . stevensoni Arnold
Petiole little more than one and a half times as one: as first tergite. Frons shining,
finely punctate. Antennal segments 10—12 about one and a half times as long as
broad; segments 4—11 with tyloidea, indistinct tyloidea on segment 12. Scutum
shining, scarcely finely punctate, prescutal sutures posteriorly slightly indicated.
Pronotal tubercles reddish-brown, sides of petiole somewhat reddish-brown. Legs
black, fore tibiae and tarsi and underside of mid and hind tibiae yellowish-brown.
Veins of wings black. No appressed silvery Pena on gaster. Length nearly 7
mmi:South;Africa vem è . . facoti sp. nov.
Scutum distinctly or ei transversely es, Une Propodeum black.
Petiole at least twice as long as first tergite. Antennae very strongly moniliform,

with tyloidea. Interantennal carina slightly or not broadened. . . . . . . 12
Scutum smooth, finely punctate, or coarsely punctate and back of propodeum with
yellow marking. Antennae not strongly moniliform . . . Licata «13

Pronotal tubercles dark brown. Fore and mid legs reddish- Bon basitarsi paler,
femora below dark brown, hind legs dark brown. Mandibles reddish. Base of petiole
dark brown or black. Gastral tergites 2—6 reddish. Pronotum and metanotum black.
Scutum with coarse punctures and light transverse rugae. Narrow smooth margin
behind propodeal enclosure, sometimes lacking. Length 8 mm. Tanzania, Rhodesia,
Kenya,rAngola, Zaire In MIE . . +. . bidentatus bidentatus (Cameron)
Pronotal tubercles yellowish or lo red. Fore and mid tibiae yellow, basitarsi
whitish. Mandibles yellowish. Base of petiole somewhat yellowish-red (not in
female). Gastral tergites 1—6 or 2—6 reddish. Pronotum partly and metanotum
more or less yellow, rarely black. Scutum with distinct punctures and feeble trans-
verse rugae. Broader smooth margin behind enclosed area of propodeum. Length 7
mm. Zaire, Equatorial Guinea, Rhodesia and Malawi . bidentatus pallidus subsp. nov.
Thorax black. Prescutal sutures long. Frontal carina more or less broadened between
antennae. Scutum finely punctate . . . ses “age A
Thorax and propodeum with yellow maine. Pu sutures na one-third
or one-half of length of scutum. Raised part of frontal carina narrow between
antennae. First gastral tergite slightly nodose. No tyloidea . . . . . . . 15
Upper part of propodeum smooth, back-side coarsely reticulate. Petiole about one
and one-third times as long as first gastral tergite, in dorsal aspect. Antennal segments
slightly rounded below, no distinct tyloidea. Pronotal tubercles yellowish-brown.

15:

16.

17.

J. P. van LITH: Revision of Ethiopian Psenini 51

Tergites 1—6 reddish. Fore and mid legs reddish, hind legs brown. Zaire .

ma benoiti Leclercq
Upper Bar of ren ie indistinctly Blans striate, back with coarse
reticulate carination. Petiole about two and one-third times as long as first tergite.
Antennal segments more cylindrical, less rounded below, segments 3—12 with very
fine narrow shining tyloidea, indistinct on twelfth segment. Pronotal tubercles dark
brown. First gastral tergite brownish, 2—6 reddish. All femora blackish-brown, fore
and mid tibiae brown. Female unknown. Zaire . . . . . . avernus Leclercq
Anterior lateral corners of pronotum obtuse. Scutum coarsely punctate, scutellum
and metanotum almost impunctate. Upper part of propodeum behind enclosed area
smooth, rest coarsely reticulate. Sides of propodeum with narrow smooth anterior
margin. Mesopleura distinctly punctate, interstices about three or four times as wide
as punctures. Petiole about one and one-third times as long as first tergite, in dorsal
aspect. Pronotum, pronotal tubercles, two square marks on scutellum, metanotum,
two marks on back of propodeum and small spot on anterior plate of mesepisternum,
brownish-yellow. Base of petiole yellowish-white, tergites 2—6 reddish, base of
second tergite black. Fore and mid legs including trochanters brownish-yellow, back
of femora brown; hind trochanters and femora brown, median part of hind tibiae
brownish, rest of hind legs reddish. Female unknown. Zaire . . trevirus Leclercq
Anterior lateral corners of pronotum sharp. Scutum shining, very finely punctate.
Upper part of propodeum smooth, back coarsely reticulate, sides of propodeum with
broad smooth anterior margin. Mesopleura finely punctate, interstices larger. Petiole
about one and a half times as long as first gastral tergite. Pronotum, pronotal tuber-
cles, two or four small longitudinal marks on scutum, scutellum, metanotum, four
marks, connected below, on back of propodeum and a small spot on anterior plate of
mesepisternum, whitish-yellow. Base of petiole yellowish-red, tergites 1—6 reddish,
first tergite more or less darkened. Fore and mid legs including trochanters pale
yellow, femora brown below; hind legs including trochanters dark brown, underside
femora and basal half of tibiae more reddish. Sierra Leone . . . oweni sp. nov.
Anterior lateral corners of pronotum with obtuse angle. Antennal segments 4—12
behind, on line between dark brown upper side and yellowish-brown underside, with
low but distinct tyloidea. Scutum with coarse variable punctation, apically finer but
more densely punctate. Upper part of propodeum behind enclosed area completely
rugoso-striate. Black; fore and mid tibiae, except brown mark below, and tarsi
yellow, base of hind tibiae whitish. Length about 7 mm. South Africa, Mozambique,
Malawi, Zambia, Tanzania, Uganda, Zaire, Sierra Leone, Ethiopia (subspecies
laevior Arnold, distinguished by brown femora and weaker sculpture of scutum and
Propodeum m. SUE : : . . « capensis capensis Brauns
Anterior lateral corners be Bro more of Te sharp, never obtuse. Antennae
without distinct tyloidea. Upper part of propodeum behind enclosed area usually
at least partly smooth, or base of hind tibiae dark. Length 7.5—8 mm . . . 17
Mid basitarsi and base of hind tibiae whitish, apart from other yellowish-white parts
of fore and mid legs. Antennae dorsally black, reddish-brown below. Punctation of
scutum distinct, interstices large, punctures usually small, rarely larger. Upper part
of propodeum behind enclosed area near median sulcus usually smooth. Anterior
lateral corners of pronotum more or less sharp. Length 8 mm. Tanzania, Rhodesia,
Uganda, Zaire, Nigeria, Sierra Leone . . Jatiannulatus latiannulatus (Cameron)

52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

— Mid basitarsi and base of hind tibiae dark brown. Antennae dorsally blackish-brown,
below on basal half reddish-brown, on apical half brown. Punctation of scutum
coarser. Upper part of propodeum behind enclosed area obliquely striate or rugose.
Anterior lateral corners of pronotum sharp. Length 7.5 mm. Zaire, Rwanda, Came-
toon | te 2 et enten ao Ane nr benleusey reel

Psen (Psen) silvaticus Arnold

Arnold, 1924: 38—39, 9 (Psen silvaticus; Rhodesia).
Leclercq, 1961a: 15, Q (Psen (Psen) silvaticus; Rhodesia).

Material studied: Rhodesia: 1 9 (holotype), “Cloudlands, 6000 ft, Vumba Mts,
6—17 April, 1923”, 1 ©, Vumba Mts, Umtali dist, 7 March, 1930, coll. R. H. R.
Stevenson, 1 ©, Vumba Mts, March, 1930, coll. R. H. R. Stevenson, 2 9, Vumba,
March, 1930, coll. R. H. R. Stevenson, 1 9, Vumba Mts, 1 March, 1938, coll. G. Arnold
(RM); 1 ©, Vumba, March, 1930, coll. R. H. R. Stevenson (SAM).

Arnold (1924) recorded the holotype as follows: “Mt. Chinzewa, 6200 ft, Vumba
Range, near Umtali, S.R. One female taken flying round a species of wild raspberry in
the hollow stems of which it probably nests. Type in R.M. coll.’ Mr. F. C. de Moor
kindly checked his topocadasteral map of the Vumba Mountains area because of the
discrepancy between the locality label under the type and Arnold’s record. In his letter,
10 May, 1973, he writes: “Cloudlands is a Private Game reserve, the highest point on
this farm is an unnamed mountain with a beacon “Castle Beacon 6245 feet”. I can find
no reference to Mt. Chinzewa however”.

Mr. de Moor also reported to me that in March, 1930 in total ten specimens were
collected in the Vumba Mts., with the following note written by Arnold: “Nesting in
cracks in old rotten tree stump (prone). All ten specimens taken from same spot in dull
weather.”

Under one of these specimens the prey has been pinned, an adult Homopteron be-
longing to the Cercopidae, about 7 mm long. Cercopidae have been recorded earlier as
the prey of Palaearctic and Indo-Australian species of Psen.

Supplementary description. — Frontal carina distinct on lower half of frons, on upper
half almost absorbed by the coarse close punctation. Arnold’s figure of the head wrongly
gives the impression that there is a transverse carina below the antennae. In reality two
very fine carinae connect the interantennal tooth with the inner side of the antennal
sclerites (Fig. 1). Propodeal enclosure ill-defined, with irregular reticulate carination,
back very finely reticulato-carinate. Hypo-epimeral area striato-punctate, like adjacent part
of mesopleura. Outer epicnemial carinae below hardly receding backwards. No acetabular
carina. Mesosternum dull, finely densely superficially punctate, fine median longitudinal
carina. Mesopleura on lower part shining, with large punctures and tendency to striation.
Upper longitudinal half of hind femora smooth, separated from normally pubescent
lower part by a narrow longitudinal area of dense fine punctation and fine short hairs,
somewhat broadened near apex, not distinctly depressed there. Petiole cylindrical with
long lateral hairs, directed obliquely downwards, sometimes worn off.

This species much resembles P. leclercgi sp. nov. from Malagasy.

J. P. vAN LITH: Revision of Ethiopian Psenini 53

Psen (Psen) congolus Leclercq

Leclercq, 1961a: 15—16, 2 and Â (Zaire).

I have examined one of the paratypes (a male) from Lubumbashi (Elisabethville),
Dec., 1933, coll. M. Bequaert (FAG). The structure of the epicnemial area and of the
mesosternum agrees with that of the Indo-Australian representatives of the subgenus
Psen. The acetabular carina is short. Hind margins of third and fourth sternites with
tufts of long brownish hairs.

Psen (Psen) leclercqi spec. nov.

Female. — Head black; mandibles reddish-brown except dark tips, labrum reddish-
brown, palpi yellowish-brown, antennae black but underside of scape, of second, third
and 12th segment reddish-brown. Thorax black, with following parts reddish-brown:
pronotum dorsally and upper part of foreside, pronotal tubercles, anterior corners of
scutum and a narrow margin along tegulae, upper two-thirds of anterior plate of
mesepisternum and tegulae. Legs black with following parts reddish-brown: fore tibiae
and tarsi, underside of mid and hind tibiae, and tarsi. Calcaria yellowish-brown. Petiole
reddish, gaster black but greater part of first and second gastral tergites, apex of last
segment and greater part of sternites reddish-brown. Veins of wings black but basal part
and stigma reddish-brown.

Clypeus feebly convex, like supra-clypeal area densely but superficially punctate,
protruding median part slightly emarginate (Fig. 2, 3). Frontal carina reaching groove
around median ocellus but almost absorbed by sculpture of frons. Carina ending below
antennae in low triangular tooth, connected with inner side of antennal sclerites by fine
carinae. Frons flat, up to ocelli densely striato-punctate, interstices shining, very narrow
margin along oculi finer and sparsely punctate. Punctation between ocelli fine, interstices
about as wide as punctures. Vertex behind ocelli with close and large punctures, laterally
coarser and somewhat striato-punctate, with only a few smooth places about as large as
punctures. Postocellar region not distinctly raised. Temples finely but distinctly punctate,
interstices about as wide as punctures or slightly more. Occipital carina ending in
hypostomal carina, both narrow and high, but lower than in P. madecassus Arnold.
Scape of antennae long, at least three times as long as wide, third segment nearly four
times, fourth segment more than twice, segments 5—7 about one and a half times, eighth
segment about one and a third times as long as wide at apex, segments 9—11 slightly
longer than wide, 12th segment about twice as long as broad at base.

Pronotal corners obtuse. Scutum densely and coarsely rugoso-punctate (Fig. 4), inter-
stices shining, medially not more than a narrow line. Prescutal sutures indicated by a
row of much finer punctures. Scutellum somewhat striato-punctate, interstices larger than
punctures, metanotum with irregular longitudinal carinae. Propodeal enclosure depressed,
lateral parts with oblique longitudinal carinae, central part with irregular carination;
back of propodeum with coarse irregular reticulate carination reaching enclosed area.
Sides of propodeum with oblique carinae and fine punctures between. Mesopleura
coarsely striato-punctate, striae posteriorly more striking, lower part finer sculptured,
hypo-epimeral area coarsely densely punctate. Epicnemial areas almost smooth, inter-
epicnemial area finely punctate, mesosternum depressed behind short acetabular carina,
antero-laterally obliquely striato-punctate, medially with strong transverse carinae. Meta-

54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

pleura shining, with some low longitudinal rugae on posterior half. Anterior plate of
mesepisternum with irregular long, partly vertical, striae; anterior oblique suture foveolate,
widened upper part with transverse striae, continuing on hypo-epimeral area. Epicnemial
carina below normally curved backwards. Metasternum with deep triangular emargination.

Femora heavy, hind tibiae dorsally with row of short thick thorns and thin long
white spines. Base of second submarginal cell of fore wings about one and a half times
as long as upper side, receiving first recurrent vein at about one-third from inner side;
second recurrent vein ending in this cell very near outer side.

Petiole as long as first two tergites and half third tergite, dorsally completely smooth,
impunctate, apically somewhat flattened, sides longitudinally slightly depressed, only
lower keel distinct, ventrally an indistinct blunt median longitudinal keel. Tergites basally
very finely punctate, on third tergite deeper than on second tergite; interstices about as
wide as punctures, on apical two-thirds interstices a few times as wide as punctures. Margin
of fourth tergite narrowly, of fifth broadly impunctate. Pygidial area elongate-triangular,
defined by distinct carina, apex rounded, surface shining but extreme apex dull, laterally
a few small punctures, each bearing a short stiff hair (Fig. 5). Sternites sparsely punctate,
somewhat more densely at base, especially antero-laterally on second sternite.

Clypeus with appressed golden pubescence and long erect hairs. Supra-clypeal area and
lower part of frons with less appressed golden pubescence; upper part of head with
longer, brownish pubescence. Temples with erect greyish pubescence, pronotum with
appressed greyish-white pubescence, thorax with long and dense pubescence, brownish
on scutum, whitish on rest of thorax, denser on mesosternum. Legs with greyish hairs,
base and apex of hind tibiae on inner side and underside of hind basitarsus with very
dense and short, yellowish-golden pubescence. Upper half of back of hind femora smooth
and shining, separated from normally pubescent lower half by a dorsally sharply defined
narrow line of fine dense punctation and short pubescence. Petiole below lower lateral
longitudinal carina with row of fine punctures, each bearing a long obliquely downwards
directed hair and also with a row of dense short hairs, densest apically. Tergites sparsely
reddish-brown pubescent, dense on lateral parts of hind margin. Apex of fifth tergite
before smooth hind margin with a few long stiff hairs, also sixth segment with long
stiff hairs. Base of second sternite with dense pale pubescence, on disk and on hind
margins of following sternites a few long stiff hairs.

Length about 11.5 mm.

Male unknown.

Malagasy: 1 ® (holotype), “Madagascar, Rogez, II.44” (FAG).

This is the second species of Psen known from Malagasy. It is easily distinguished
from P. madecassus Arnold, which is smaller, more reddish, and much more finely, very
sparsely, punctate, especially on front, vertex, scutum and sides of thorax.

P. leclercqi is very similar to P. silvaticus Arnold from Rhodesia but it differs not only
in the reddish anterior corners of the scutum, but especially in the sculpture of the
propodeum. In P. si/vaticus the enclosed area is hardly distinguishable, with irregular
reticulate carination; back-side of propodeum very finely reticulato-carinate. In P. leclercgi
the propodeal enclosure is well-defined, lateral parts of enclosure shining, with oblique
distinct carinae, median part with coarse irregular sculpture, back of propodeum
coarsely irregularly reticulato-carinate.

J. P. van LITH: Revision of Ethiopian Psenini 55

Fig. 1. Psen (Psen) silvaticus Arnold, @. Face. Fig. 2—5. Psen (Psen) leclercqi sp. nov., 2, holo-
type. 2, face; 3, clypeal margin; 4, punctation median part of scutum; 5, pygidial area

56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

Psen (subgenus?) madecassus Arnold

Arnold, 1945: 152—153, $ and & (Psen madecassus; Malagasy: Bekily, Ranomafana, Ivondro).
Leclercq, 1961a: 15, $ and 4 (Psen (Psen) madecassus; Malagasy: Bekily, Rogez).
Leclercq, 1961c: 117, 2 (Malagasy: Mandraka).

Material studied: Malagasy: 1 ©, paratype and 1 G', paratype, both labelled “in cop.
B.”, Bekily, Febr., 1937, coll. A. Seyrig, 2 &, Bekily, Jan., 1937, coll. A. Seyrig, 1 9,
Perinet, “Forêt côte est”, Febr., 1939, coll. A. Seyrig (RM); 1 9, Bekily, Jan., 1942
(BM); 1 &, Bekily, Jan., 1937, coll. A. Seyrig (author's collection); 1 g', Rogez, June,
1944, coll. A. Seyrig (MRAC); 1 9, Madagascar Est, Mandraka (Tamatave), 4 April,
1958, coll. F. Keiser (NMB).

The excellent description by Arnold may be supplemented with the following char-
acters. — Frontal carina fine, reaching anterior ocellus, slightly widening at lower end
and ending in a low small tubercle. Lateral parts of frons with large low tubercle. In
both sexes occipital carina narrow and high, especially lower part, ending normally in
hypostomal carina. Apical margin of clypeus thick, slightly emarginate.

Scutum coarsely punctate, interstices as large as, sometimes larger than punctures.
Mesopleura not closely and finely punctate, as stated by Arnold, but finely and sparsely,
including hypo-epimeral area, interstices many times larger than punctures. Mesosternum
depressed and smooth behind interepicnemial area, with high median longitudinal carina,
anteriorly defined by a distinct short acetabular carina. Metasternum deeply emarginate.
Petiole dorsally smooth, latero-dorsally with shallow narrow groove, sides of petiole with
distinct groove, also ventro-laterally a distinct longitudinal groove; in all these grooves a
row of fine punctures, each with a long erect fine hair. Hind coxae with sharp inner longi-
tudinal carinae. Upper third of hind femora smooth, medially a broad, densely finely
punctate and pubescent, longitudinal line, lower part of femora sparsely longer pubescent.
Pygidial area of female (Fig. 13) elongate-triangular, apex emarginate, lateral carinae
high. Antennae of male without distinct tyloidea. No fasciculate hairs on apical margins
of third and fourth gastral sternites, but margins of second and following sternites with
fine pale, obliquely inwards directed, long erect hairs. Genitalia of male: Fig. 6, 7, 8, 9.
Seventh sternite: Fig. 10. Eighth sternite: Fig. 11, 12.

P. madecassus does not belong to the subgenus Pser because of the somewhat different
shape of the sternum, the fine, punctate, latero-dorsal sutures of the petiole and the
absence of fasciculate hairs on the apical margins of the third and fourth gastral sternites
of the male. Probably it should best be placed into the subgenus Punctipsen, of which
representatives are known from the Palaearctic region, Ryukyu Islands, Formosa and S
India. The characters they have in common are the punctation of the petiole, the almost
vertical foreside of the first tergite, the continued longitudinal rugae on the back of the
propodeum, behind the enclosed area, the thick apical margin of the clypeus, the absence
of fasciculate hairs on the sternites of the male and to some extent also the shape of the
male genitalia.

Fig. 6—12. Psen madecassus Arnold, &. 6, outer valve of genitalia, dorsal aspect; 7, the same,

lateral aspect; 8, the same, inner aspect, with volsella; 9, right penis valve, outer side; 10, 7th

sternite, ventral aspect; 11—12, 8th sternite, lateral and ventral aspect. Fig. 13. Psen madecassus

Arnold, 9, pygidial area. Fig. 14—18. Psen patellatus Arnold, 6, paratype. 14, face; 15, 6th, 7th
and 8th sternites, ventral aspect; 16—18, genitalia, ventral, lateral and posterior aspect

J. P. van LITH: Revision of Ethiopian Psenini

58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

P. madecassus differs from the earlier described species belonging to Punctipsen in the
reduced punctation of the dorsal side of the petiole and in having no tooth on mandibles
or mid basitarsi of the males, but I think that these characters do not have any sub-
generic value.

The hind coxae have a sharp inner carina as in the subgenus Pseneo, but there is no
oval patch on the posterior apex of the hind femora, the clypeus is bidentate and the
petiole has latero-dorsal grooves with fine punctures and long erect lateral and ventral
hairs.

Arnold (1945) records one specimen with prey, a small adult Cercopid.

Psen (subgenus?) patellatus Arnold

Arnold, 1924: 39—40, & (Psen patellatus; SW Africa: Onoolonga and Mafa).
Leclercq, 1961a: 15 (Psen (Psen) patellatus)).

Material studied: South-West Africa: 1 & (holotype), Mafa, Febr., 1923, 1 {,
Onoolonga, Febr., 1923, det. Arnold (SAM); 1 g', Onoolonga, Febr., 1923, with green
paratype label (RM).

Dr. Hesse of the South African Museum informed me that there are two more males
in his collection, respectively one from Mafa and one from Onoolonga. Arnold (1924)
recorded in total three males from Onoolonga and Mafa, he did not mention which was
the holotype.

Little can be added to the detailed original description. — Face broad (Fig. 14),
hypostomal carina ending normally in occipital carina. Mandibles unidentate at apex.
Sternum very densely punctate, no acetabular carina, lateral epicnemial carinae not curved
backwards as in subgenus Psen. Petiole short, not only dorsally, but also laterally with
irregular longitudinal carinae; similar carinae on ventral side which are converging back-
wards; ventral plate with fine longitudinal rugae. Fourth and fifth gastral sternites with-
out fasciculate hairs. Sternites 6—8: Fig. 15, ventral aspect. Apical spine short. Genitalia
short, base and apex dark brown, median part and penis valves testaceous (Fig. 16, 17
and 18).

Arnold already doubted whether this species belongs to the subgenus Psen. The
structure of the epicnemial areas and of the petiole, the depressed posterior side of the
propodeum, the absence of fasciculate hairs on the apical margins of the fourth and fifth
sternites, and the short genitalia make it very unlikely that P. patellatus should be placed
in the subgenus Psen. As it does not fit in any of the other subgenera, perhaps a new
subgenus has to be erected to receive this species.

Psen (Mimesa) empeyi spec. nov.

Male. — Head and thorax black; anterior margin of lower part of pronotum somewhat
thicker and yellowish-white; apices of mandibles reddish, palpi very dark brown, antennal
segments 5—12 below, last two segments completely, orange-reddish, segments 5—11
brownish dorsally. Tarsi brownish, tibial spurs of hind legs whitish. Veins of wings
brown, base of veins and base of stigmata of fore wings paler brown. Petiole black,
ventral plate red, also first tergite and base of second tergite, following tergites black.

No distinct frontal carina, small low tubercle between antennae. Frons and vertex
densely punctate, interstices on vertex larger, at least as wide as diameter of punctures,

J. P. vAN LITH: Revision of Ethiopian Psenini 59

shining. Temples rather densely punctate, surface with fine striae. Underside of head shin-
ing, occipital carina distinctly ending in hypostomal carina. Face below antennae, including
clypeus, dull, finely very densely punctate, protruding median part of clypeus with small
triangular emargination and slight depression (Fig. 19). Mandibles narrow. Third
antennal segment nearly three times, fourth segment about twice, fifth segment about one
and a half times, segments 6—8 about one and a quarter times as long as broad at apex,
segments 9—12 about as long as broad at apex, last segment about one and three-quarter
times as long as wide at base. Interocellar area almost impunctate, distance between
posterior ocelli longer than distance between ocelli and oculi.

Sides of pronotum converging roundly backwards (Fig. 20). Scutum shining, densely
punctate, interstices mostly larger than diameter of punctures, prescutal sutures indistinct,
scutellum punctate as scutum, posteriorly somewhat longitudinally striate. Metanotum
dull, with coarse and very close reticulate carination. Propodeum also dull, with coarser
reticulate carination, coarsest on lower half. Propodeal enclosure ill-defined, propodeal
longitudinal groove deep, coarsely reticulato-carinate. Sides of propodeum dull, finely
closely reticulato-carinate, more striate on anterior half. Upper part of metapleura dull,
lower part almost shining. Mesopleura, anterior plate of mesepisternum, greater part of
epicnemial areas and mesosternum densely punctate, with narrow shining interstices;
hypo-epimeral area dull, finely closely reticulato-carinate or coarsely coriaceous, epimeral
suture ill-defined. First recurrent vein of fore wings interstitial, second recurrent vein
ending near end of second submarginal cell. Legs slender. Petiole about one and a half
times as long as first tergite, thin, cylindrical, slightly widened at base and apex, with
small pit dorsally at apex. Gaster slender, finely alutaceous, last tergite dorsally flattened,
broad-triangular with rounded apex (Fig. 21, 22), densely punctate with short appressed
hairs.

Sixth sternite dentate apically (Fig. 23). Seventh sternite whitish transparent, tip dark
brown with triangular emargination (Fig. 24). Eighth sternite whitish transparent, apical
margin dark brown, spine short and dark brown (Fig. 25, 26). Genitalia dark brown:
Fie227,28.

Pubescence silvery-whitish, dense and appressed below antennae.

Length about 6.5 mm.

Female unknown.

South Africa: 3 & (holotype and paratypes), Ellisras, N. Transvaal, 19 June, 1962,
1 & (paratype), 22 June, 1963, coll. H. N. Empey (holotype deposited by Mr. Empey
in the Transvaal Museum, paratypes EC, PMFV and in the author's collection).

P. empeyi is the first known Ethiopian species of the subgenus Mimesa, a subgenus
well represented in the arctic and Mediterranean regions. It is distinguished by the
remarkable female-like pygidial area which resembles that of the Nearctic Psen (Mimesa)
pygidialis Malloch (1933).

Psenulus turneri Arnold

Arnold, 1927: 121, 2 (South Africa: Cape Province).
Leclercq, 1961a: 26.

Supplementary description of female. — Outer half of mandibles reddish. Fourth
tergite varying from completely reddish to basally or almost completely black. All of the
eleven females from Grahamstown and Kenton-on-Sea, mentioned below, have darker

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

J. P. VAN LITH: Revision of Ethiopian Psenint 61

legs than the holotype, the hind legs are completely black. The holotype has a completely
black fourth tergite and all legs are reddish.

Frontal carina raised and much broadened between antennae, connected with high
transverse carina below antennae by two closely parallel or confluent carinae. Prescutal
sutures long, on anterior three quarters of scutum with short transverse carinae (alveoles).
Petiole cylindrical, little longer than first tergite in dorsal aspect. No distinct pygidial
area. One of the six females from Howison’s Poort, Grahamstown, has a triangular,
instead of petiolate, second submarginal cell, in both wings.

Apical margin of fourth and fifth gastral sternites with dense fringe of long whitish
hairs.

First description of male. — Resembling female. Mandibles almost completely reddish.
Basal half of fourth tergite black in all three specimens. Hind legs partly brownish.
Antennal segments 3—13 brown above; scape, second segment and underside of flagel-
lum reddish.

Antennae moniliform, segments 3—8 about one and a half times, segments 9—12
about one and a third times as long as broad at apex, last segment about one and three
quarters times as long as broad at base, segments 4—8 with small narrow tyloidea,
segments 9—10 with indistinct tyloidea. Petiole of second submarginal cell shorter than
in female. Back of propodeum coarsely reticulate, more or less striate behind enclosed
area. Petiole slightly shorter than in female, below with distinct lateral keel. No conspic-
uous fringes on apical margins of fourth and fifth gastral sternites.

Apex of seventh sternite with triangular emargination (Fig. 31) and fringe of long
fine hairs. Eighth sternite: Fig. 29, 30. Genital apparatus (Fig. 32, 33) short, broad,
pale yellowish-brown, stipes broad and rounded at apex.

The holotype is a female from South Africa, Cape Province, Mossel Bay, 18—30 Nov.,
1921, coll. R. E. Turner (BM).

New records from South Africa, Cape Province: 6 9, Howison’s Poort, Grahamstown,
19—30 Nov. and 14—20 Dec., 1971, 2 9, Belmont Valley, Grahamstown, 28—31
Oct. and 5—9 Nov., 1971, all collected by F. W. Gess, 3 9, Kenton-on-Sea, 1—9 Nov.,
1971, 26-31 Nov. 1971 and 1—14 March, 1972, coll. R. A. Jubb; 1 g', Howison’s
Poort, Grahamstown, 24—26 Nov., 1971, coll. F. W. Gess, 3 d, Kenton-on-Sea, 23—
29 Oct., 1970, 14—21 Oct., 1971, 21—26 Oct., 1971, coll. R. A. Jubb, all collected in
Malaise traps (AMG).

Leclercq (1961a) rightly assumed that P. twrneri and P. paultsae Leclercq are very
closely related, but they are still distinctly different.

Psenulus paulisae Leclercq
Leclercq, 1961a: 23—26, 2 and & (Zaire).

New records from Sierra Leone: 1 9, Njala, Dec., 1934, coll. E. Hargreaves, pres. by
Com. Inst. Ent. B.M. 1973—1 (BM); Freetown, 4 ©, March, May, July, 1967, 3 3,
Febr., April, 1968, coll. D. F. Owen (HT).

Fig. 19—28. Psen (Mimesa) empeyi sp. nov., 6, 19, holotype, 20—28, paratype. 19, face; 20,

pronotum; 21—22, 7th segment, dorsal and latero-dorsal aspect; 23, apex 6th sternite, dorsal aspect;

24, 7th sternite, latero-ventral aspect; 25—26, 8th sternite, ventral and lateral aspect; 27, genitalia,

dorsal aspect; 28, apex penis valve, lateral aspect. Fig. 29-33. Psenulus turneri Arnold, &. 29—30,

Sth sternite, with part of musculature, ventral and lateral aspect; 31, 7th sternite, lateral aspect;
32—33, genitalia, right stipes bent, dorsal and lateral aspect

62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

1 have compared these specimens with a paratype (female) from the type-locality
Paulis (FAG). In the females the transverse striae on the sides of the scutum are less
distinct, but in one of the males they are distinct. Leclercq has given a very good and
detailed description but I would like to mention the following additional characteristics.

Female. — Anterior oblique suture of mesopleura foveolate, widened upper part with
transverse carinae. Stiff hairs of fringes on apical margins of fourth and fifth sternites
about as long as fourth segment of posterior tarsus.

Supplementary description of male based on the males from Sierra Leone. — Antennal
segments moniliform. Scape, second segment and underside of following segments
yellowish-red, dorsal side of flagellum dark brown, third, 12th and 13th segment some-
what paler. Third antennal segment about one and a half times as long as broad at apex,
length of segments then gradually decreasing, segments 11—12 about as long as broad
at apex, last segment about one and a half times as long as broad at base. Segments 3—9
with distinct, narrow and oblique, pale tyloidea, on tenth segment much smaller. Raised
part of carina between antennae somewhat narrower than in female.

There is some variation in the colour of the gaster. One of the females from Sierra
Leone, collected March, 1967, has also the first tergite completely reddish, and the apex
of the ventral plate of the petiole. In one of the males from Sierra Leone not only the
first tergite and at least the base of the second tergite are deep black, but the following
tergites also are somewhat darkened on the sides. Apical margins of sternites 2—6
brownish. Also the sculpture of scutum and propodeum may vary somewhat.

Psenulus erusus Leclercq

Leclercq, 1961a: 26—27, Q (Zaire).

New records: Sierra Leone: Freetown, 1 ©, July, 1967, 2 9, Jan., 1968, coll. D. F.
Owen (HT).

These specimens are identical with the holotype from Zaire (Congo), Eala, Nov., 1936
(MRAC), but they are slightly smaller and the upper part of the back of the propodeum
is less densely striate.

Length about 6.5 mm.

There are fringes of long hairs on the apical margins of the fourth and fifth sternites
like in the other species belonging to the group of P. turneri.

Psenulus garambae Leclercq

Leclercq, 1961b: 57—58, 9 (Zaire).

In the holotype the dilated raised part of the interantennal carina is wide, ratio width-
length being about 1 : 2.5, carina ending below antennae in a distinct transverse carina.
Tempora coarsely striato-reticulate. Petiole about one and one-third times as long as first
gastral tergite.

Apical margins of fourth and fifth sternites with fringe of long, yellowish golden
hairs.

Length about 6 mm.

Leclercq recorded this specimen from the National Park of Garamba, in wooded
savanna surrounding the camp of Bagbele, 20 March, 1950, coll. H. de Saeger (MRAC).
There are no further records. He has already recognized the close relationships of P.
garambae with P. saltitans Arnold, as well as with P. turneri, P. paulisae, and P. erusus.

J. P. vAN LITH: Revision of Ethiopian Psenint 63

Psenulus patei Arnold

Arnold, 1940: 132—133, £ (Rhodesia: Bulawayo and Khami).
Leclercq, 1961a: 29, 9 (Rhodesia: Matopos).

Arnold based his description of this species on 6 females from Bulawayo and Khami.
The holotype, which Mr. F. C. de Moor of the National Museum at Bulawayo kindly sent
me on loan, is labelled: “Khami, S. Rhodesia, 30.10.1938, Nat. Museum S. Rhodesia”
and also bears a red type-label of G. Arnold.

The remaining five females recorded by Arnold are undoubtedly included in the
following material from the type localities: 2 9, Bulawayo “Kh” (= Khami?), 16
Sept., 1928, 1 9, Bulawayo, 25 Dec., 1929, 1 9, Khami Dam, 14 Sept., 1938, 2 9,
Khami, 1 Nov., 1938, all collected by R. H. R. Stevenson (RM).

I could also study the female recorded by Leclercq (1961a) from Rhodesia: Matopos,
20 Nov., 1938, leg. et det. G. Arnold (BM).

New records from Rhodesia: 1 9, Murampara, 9 Sept., 1927, 1 9, Umguza Valley,
Bulawayo dist., 1 Febr., 1930, both coll. R. H. R. Stevenson (RM).

Supplementary description, based on holotype. — Mandibles dark reddish except
blackish tips. Labrum dark reddish. Palpi pale brown. Fore legs yellowish-red, trochanters
completely and base of femora below brown, basitarsi yellowish-white. Mid trochanters
and femora brown, apex of mid femora yellowish-red below, base of mid tibiae yellowish,
underside pale reddish-brown. Hind trochanters, femora and tibiae brownish-black, basal
fourth of tibiae straw-yellow, base of hind basitarsi pale yellowish. Pronotal tubercles
dark brown, tegulae yellowish-transparent. Veins of wings dark brown.

Frons below antennae with tendency to transverse striation. Vertex shining, very finely
punctate. Labrum not distinctly tridentate. Antennae short, third segment about one and a
half times as long as broad at apex, segments 4—6 little longer than broad, segments
7—11 shorter than broad at apex, last segment about one and a half times as long as
broad at base.

Prescutal sutures almost reaching hind margin of scutum, indicated only by a row of
irregular short transverse grooves. No pygidial area. Apical margins of fourth and fifth
sternites with fringes of long yellowish-golden hairs, sixth sternite densely yellowish-
golden pubescent. Epicnemial areas below with patch of dense short whitish pubescence.

The female from Umguza Valley has darker mandibles, reddish fore tarsi, light brown
mid tibiae, straw-yellowish mid basitarsi, rest of mid tarsi pale reddish, brownish-black
hind legs and yellowish-brown base of hind tibiae.

P. patei is recognized by the triangular (rarely slightly open) second submarginal cell
of the fore wings, striate upper part of propodeum and reddish tergites 4—6. It is closely
related to P. saltitans Arnold, which also belongs to the group of P. turner? Arnold but
P. saltitans has no distinct pale base of hind tibiae.

According to Arnold (1940) this species was found nesting in a decayed tree trunk.

Psenulus saltitans Arnold
Arnold, 1958: 135, 9 (Rhodesia).
Leclercq, 1961a: 29.
Material studied: 1 Q, “Cashel, S. Rhodesia”, 30 Dec., 1947, with green paratype-
label (RM).
Supplementary description. — Middle part of mandibles dark reddish. Fore trochanters

64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

and at least outer side of fore femora brown, not reddish-brown. Mid basitarsi straw-
yellow. Veins of wings brown.

Frons shining, oblique rugae superficial. Antennae short, third segment about one and
one-quarter times as long as broad at apex, segments 4--6 little longer than broad,
segments 7—11 wider than long, last segment about one and three-quarter times as long
as broad at base. Interantennal carina raised and broadened between antennae, lateral
edges of widened part converging below and ending as two very close parallel carinae in
the transverse carina below antennae. Transverse carina more angular than figured by
Arnold (1958: 134). Lower half of clypeus shining, impunctate.

Prescutal sutures as in P. patei Arnold. Anterior oblique suture foveolate, widened
upper part with transverse carinae reaching upper anterior corner of episternal area. No
pygidial area. Petiole about one and a half times as long as first tergite in dorsal aspect,
cylindrical, base flattened dorsally.

Petiole dorsally with lateral row of very short hairs, ventro-laterally a few long out-
standing hairs. Apical margins of fourth and fifth sternites with fringe of long yellowish-
golden hairs, sixth sternite yellowish-golden pubescent.

P. saltitans belongs to the group of P. turneri Arnold, but differs from latter species in
the non-petiolate but triangular second submarginal cell of the fore wings. It differs from
P. patei in the dark bases of mid and hind tibiae.

According to Arnold this species was caught “jumping about on the leaves of a shrub,
probably in search of aphids”.

Psenulus ealae Leclercq

Leclercq, 1961a: 27—29, 9 (Zaire).

New records: Sierra Leone: Freetown, 1 9, May, 1967, 1 g', April, 1968, coll. D. F.
Owen (HT).

A series of seven females is known from Eala; through the kind intermediary of Prof.
J. Leclercq I could study three paratypes (FAG).

Supplementary description of female based on these paratypes. — All legs brown,
apex of fore femora, fore tibiae, and underside of mid tibiae more yellowish-brown, base
of hind tibiae yellowish-grey. Veins of fore wings dark brown.

Transverse carina below antennae not long, reversed-V-shaped with an angle of about
100 degrees. Foreside of pronotum with transverse striae. Anterior oblique suture broad,
foveolate, widened upper part with transverse carinae. Base of second sternite with semi-
circular depression. No pygidial area. Apical margins of fourth and fifth sternites with
fringe of long yellowish-golden hairs.

Length about 6.5 mm.

First description of male (Sierra Leone). — Resembling female. Base of fourth tergite
also brown. Raised and broadened part of interantennal carina narrower, ratio width-
length about 1:4. Antennal segments convex below, third segment about one and
three-quarter times, segments 4—12 about one and a half times as long as broad at apex,
last segment nearly two and a half times as long as broad at base. Segments 4—10 behind
with distinct narrow oblique tyloidea, on segments 5—6 about half as long as segment,
all reddish like underside of flagellum, on segments 3 and 11 at most a small point.

Second submarginal cell of fore wings sessile, but its upper width shorter than in

J. P. van LITH: Revision of Ethiopian Psenini 65

Fig. 34—36. Psenulus aurifasciatus sp. nov. 2, holotype. 34, face; 35, frontal (interantennal)

carina, dorsal aspect; 36, propodeal enclosure. Fig. 37—41. Psenulus freetownensis sp. nov, 9,

holotype. 37, face; 38, frontal carina, dorsal aspect (39, the same, paratype); 40, 2nd and 3rd sub-
marginal cell of left fore wing; 41, propodeal enclosure

female. Petiole about one and a half times as long as first tergite. This is much longer
than in the female, in which the petiole is about equal in length. However, this may be
a sexual difference.

The female from Sierra Leone is somewhat smaller than the paratypes from Eala, the
scutum is slightly less strongly sculptured and the fore femora are more reddish.

P. ealae is closely related to P. turner? Arnold, but is distinguished easily by the open
second submarginal cell.

Psenulus aurifasciatus spec. nov.

Female. — Head and thorax black; mandibles partly brownish, palpi yellowish-brown.
Underside of antennae reddish-brown. Apex of fore femora, fore tibiae completely and
tarsi reddish; back-side of mid tibiae and tarsi reddish, rest of mid legs and hind legs
completely black or very dark brown. Gaster black, apical margins of tergites 2—5
brownish transparent laterally; last tergite, greater part of fifth and whole of sixth sternite
brown. Veins of wings dark brown, tegulae reddish-brown.

66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

Frontal carina (Fig. 35) raised and broadened between antennae, ratio width-length
about 1 : 2.5, lower narrow part of carina about as long as broadened part and ending
below in a transverse carina, its length about half the distance between the eyes, obtusely
angular in the middle (Fig. 34). Basal half of clypeus dull, apical half shining, median
part much protruding with low triangular emargination, forming two triangular teeth.
Mandibles normal. Occipital carina ending in hypostomal carina. Frons near oculi with
large convex, shining and very finely punctate areas, below anterior ocellus on both sides
of carina very densely finely punctate. Vertex shining, very finely punctate, punctures
between ocelli coarser. Tempora with coarse, striato-reticulate longitudinal carination,
especially along the eyes. Antennae short, third segment about one and a half times,
segments 4—5 about one and one-third times as long as broad at apex, segments 6—10
almost square, 11th segment about one and one-quarter times as long as broad at apex,
last segment about twice as long as broad at base.

Pronotal corners rounded, foreside above with fine longitudinal striation. Scutum
shining, finely punctate, prescutal sutures linear on anterior fourth, on posterior part
until near hind margin of scutum indicated by a line of punctures larger than those on
rest of scutum. Scutellum and metanotum very finely punctate, interspaces a few times
as large as punctures. Lateral parts of propodeal enclosure medially separated from
metanotum by two triangular, dull areas; median part broad, almost heart-shaped and
connected with anterior margin by a small separate area (Fig. 36). Back of propodeum
shining, dorsally with superficial oblique striae, followed by a more coarsely and some-
what irregularly striate area, lower third with fine irregular reticulation and a few
stronger carinae. Median longitudinal groove broad. Posterior half of sides of propodeum
reticulate, separated from back-side by a broad, dorsolateral, irregularly foveolate groove;
anterior part of sides dull. Metapleura dull over greater part. Mesopleura and mesoster-
num shining, distinctly punctate, interstices mostly larger than punctures, punctures very
fine on hypo-epimeral area, a foveolate groove along metapleura. Anterior oblique suture
broad and foveolate, widened upper part with transverse carinae. Legs normal, stoutly
built, a row of four small teeth on apical two-fifths of mid tibiae, hind tibiae with two
very small thorns on apical half. Second submarginal cell of fore wings sessile, upper
side about one-fifth of length of lower side, first recurrent vein ending in second sub-
marginal cell. Petiole about one and one-quarter times as long as first tergite, cylindrical,
base above distinctly, and sides only slightly, flattened, apex above with a small pit.
Gaster very finely punctate. No pygidial area. Second sternite at base with a rounded-
triangular depression.

Face with appressed silvery pubescence, sparse on anterior third; also with long whitish
hairs. Head, thorax and legs whitish pubescent, more yellowish-grey and longer on vertex,
dense and somewhat yellowish-grey on lower part of epicnemial areas and on mesoster-
num. Gaster with yellowish-brown pubescence. Pygidium with many long stiff hairs,
apical margins of fourth and fifth gastral sternites with fringe of long, yellowish-golden
hairs.

Length 8 mm.

Male unknown.

Sierra Leone: 1 9 (holotype), Freetown, April, 1967, coll. D. F. Owen (HT).

This species belongs to the group of P. turneri and is easily distinguished by its dark
gaster.

J. P. vAN LITH: Revision of Ethiopian Psenini 67

Psenulus freetownensis spec. nov.

Female. — Head and thorax black; mandibles reddish, palpi testaceous. Scape of
antennae and underside of flagellum yellowish-brown, flagellum above dark-brown.
Pronotal tubercles and tegulae yellowish-brown. Veins of wings dark-brown. Legs in-
cluding trochanters reddish, fore and mid legs slightly paler than hind legs. Petiole,
except ventral plate, black, following segments of gaster reddish.

rontal carina (Fig. 38, 39) raised and widened between antennae, length of widened
part about 3.5 times width, carina ending below antennae in transverse carina, which has
the shape of a reversed V with an angle of about 120 degrees (Fig. 37). Clypeus very
finely and densely punctate, lower margin less densely punctate and more shining;
protruding median part with two distinct small teeth, distance between apices of teeth
about one sixth of total distance there between eyes. Frons and vertex shining, distinctly
punctate. Tempora with coarse reticulato-striate sculpture behind eyes. Occipital carina
ending normally in hypostomal carina. Mandibles normal. Third antennal segment at
least one and a half times, segments 4—5 about one and a third times as long as broad
at apex, segments 6—7 slightly longer than broad, segments 8—11 about as long as
broad at apex, last segment about one and three quarters times as long as broad at base.

Lateral corners of pronotum obtuse. Scutum densely finely punctate, prescutal sutures
not sharp, almost reaching hind margin, parapsidal sutures distinct. Scutellum and meta-
notum very finely and densely punctate. Propodeal enclosure triangular, central part
broad, separated from metanotum by a small square area (Fig. 41), lateral areas with
at least three oblique carinae. Median longitudinal groove broad, oval, with narrow
deeper groove. Upper half of back of propodeum smooth and shining, lower half with
fine reticulate carination. Posterior half of sides of propodeum with reticulate carination,
anterior half opaque, propodeum postero-laterally with a broad foveolate groove. Meta-
pleura opaque, somewhat coriaceous, mesopleura and mesosternum shining, very finely
punctate. Anterior oblique suture broad, foveolate, broadened upper part with transverse
carinae. Upper side of second submarginal cell about one third length of lower side of
cell, first recurrent vein ending just in second submarginal cell, second recurrent vein
ending well in third submarginal cell (Fig. 40). Legs normal, mid tibiae at apex behind
with 3—5 short reddish thorns, on back of hind tibiae in the middle two or three short
reddish thorns.

Petiole about two-thirds of length of first tergite (in dorsal view), almost cylindrical,
base flattened and with lateral carina, apex with small pit. Gaster very finely punctate.
No pygidial area. Second sternite at base with semi-circular depression.

Face with silvery appressed pubescence and also with long erect silvery hairs. Pubes-
cence of head and thorax whitish, epicnemial areas below with round patch of dense
appressed pubescence. Petiole with long lateral hairs and dorsally with two backwardly
convergent rows of very short hairs. Pubescence of legs and gaster yellowish-golden,
apical margins of fourth and fifth sternites with fringe of long, yellowish-golden hairs,
sixth sternite with short dense yellowish-golden pubescence.

Length about 6.5—7 mm.

Male. — As female, but underside of antennae paler, transverse carina below antennae
longer and less angular in frontal aspect, segments 3—12 about one and a half times as
long as broad at apex, last segment about two and a half times as long as broad at base,
underside of segments convex, segments 4—8 on back with indistinct, oblique, reddish

68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

tyloidea. Circular patches of appressed pubescence at lower side of epicnemial areas less
conspicuous and sternites 4—5 without fringes of long hairs. Upper, horizontal part of
propodeum smooth and shining, back and sides of propodeum more coarsely sculptured
than in female. Petiole on apical half below with distinct ridge, sides of petiole slightly
depressed.

Length about 5.5 mm.

Sierra Leone: Freetown, 1 9 (holotype), Aug., 1967, 1 g' (allotype), July, 1967,
1 & (paratype), Jan., 1968, all collected by D. F. Owen (HT, paratype in author's
collection).

Equatorial Guinea (Spanish Guinea): 1 9 (paratype), ‘“Uelleburg”, June—Aug.,
1908, coll. Tessmann (ZMB).

In the allotype the second submarginal cell of the left wing is almost triangular. On
the contrary this cell is more open above in the male paratype than in the holotype.

The petiole of the female from Equatorial Guinea is brownish, its apex reddish
dorsally. The interantennal carina is slightly broader and the upper side of the second
submarginal cell is about one fourth of the length of the lower side.

P. freetownensis is in both sexes easily recognized by the completely red gaster and
legs, and the broad smooth upper part of the propodeum behind the enclosed area. It
evidently belongs to the group of P. turneri, of which some species normally have triangu-
lar or even petiolate second submarginal cells.

Psenulus alveolatus spec. nov.

Male. — Head and thorax black; antennae brown, underside and scape completely
reddish-brown. Palpi pale brown. Tegulae dark reddish-brown. Labium and apical half
of mandibles reddish-brown. Fore legs including trochanters reddish-brown, mid and
hind legs brown. Tibial spurs of mid and hind legs yellowish-white. Petiole including
ventral plate deep black, also first tergite and basal two-thirds of second tergite, rest of
dorsal side of gaster reddish; sternites 2—4 very dark brown, nearly black, following
sternites reddish; apical spine dark reddish. Veins of wings dark brown.

Frontal carina raised and much broadened between antennae, ending below antennae
in high transverse carina. Clypeal margin medially with two low triangular teeth. Frons
with coarse oblique rugae, interocellar area with transverse rugae, vertex more shining,
striato-punctate, striae weak. Tempora coarsely striato-reticulate with a few strong longi-
tudinal carinae. Occipital carina ending in hypostomal carina, high, alveolate along fore-
side. Antennal segments 3—8 about one and a half times, segments 9—12, about one and
a quarter times width at apex, last segment one and three quarters times width at base;
third segment with small tyloidea, segments 4—8 with longer and distinct, narrow
tyloidea.

Anterior margin of pronotum alveolate; scutum shining, with irregular rather dense
punctation, and centrally with weak transverse rugae; prescutal sutures with deep and
large alveoles and strong transverse rugae, reaching posterior margin which shows many
short longitudinal rugae; parapsidal sutures distinct; lateral margins of scutum alveolate.
Scutellum shining, sparsely finely punctate. Metanotum somewhat more densely punctate.
Enclosed area of propodeum shining, with rather short oblique carina on lateral parts,
wide central area; back coarsely reticulate, median longitudinal carina deep and wide but
irregular. Sides of propodeum less coarsely reticulate. Metapleura smooth. Mesopleura

J. P. vAN LITH: Revision of Ethiopian Psenini 69

and mesosternum shining, densely coarsely punctate, partly somewhat rugose. Anterior
plate of mesepisternum shining, less densely punctate, not rugose. Anterior oblique
suture broad, foveolate, widened upper part with transverse carinae.

Legs slender. Second submarginal cell distinctly petiolate, first recurrent vein ending
in second submarginal cell, second recurrent vein ending well in third submarginal cell.
Petiole long, nearly twice length first tergite; sides slightly depressed, with distinct lower
keel, dorsally convex, but base and apex somewhat flattened, narrow triangular pit at
apex.

Face with appressed silvery pubescence and erect longer hairs, rest of head, and thorax,
with whitish hairs, gaster yellowish-grey pubescent, apical margins of sternites with thin
fringe of short, backwards directed stiff hairs, disk of sixth sternite with dense, back-
wards directed, golden pubescence.

Length about 7 mm.

Female unknown.

Rhodesia: 1 & (holotype), Bulawayo, 25 Dec., 1929, coll. R. H. R. Stevenson (RM).

P. alveolatus is certainly closely related to P. twrneri Arnold, differing however by
having a longer petiole, a more coarsely sculptured scutum, and by the fact that more
gastral segments are reddish.

A male from Angola (A30), 7 miles W. of Gabela, 16—18 March, 1972, Southern
African Exp. B.M. 1972—1 (BM) is very similar and certainly belongs to the group of
P. turneri. It has also a long petiole and tergites 2—6 are completely reddish, but it
differs from alveolatus in being somewhat smaller, with the scutum less coarsely sculp-
tured and the second submarginal cell being open dorsally. As the antennae are missing
I prefer to wait for additional material before describing this form as a new species.

Psenulus fulgidus Arnold

Arnold, 1945: 156—157, ® and 4 (Malagasy: Bekily, Behara and Ranomafana).
Leclercq, 1961a: 35, 9 (Malagasy: Fort-Dauphin).

Material studied: 2 9, Bekily, Dec., 1932, Febr., 1940, 1 &, Bekily, May, 1934,
(FAG); 1 9, Bekily, March, 1937, coll. A. Seyrig, det. Arnold (?) (author’s collection).

The material from the collections of the Faculty of Agricultural Sciences at Gembloux
was kindly sent to me by Prof. J. Leclercq; it forms part of the material collected by
Seyrig and was mounted after Arnold’s study (1945) and identified by Leclercq (1960).

P. fulgidus and P. reticulosus Arnold are much alike as regards their colour patterns,
but they differ in a few important characteristics. P. fulgidus is smaller (female 6.3—8.5
mm, male 6—6.5 mm, cf. Arnold, 1945, who based his description on 16 females and 9
males). In the female of P. fulgidus the broadened raised part of the interantennal carina
is lozenge-shaped, petiole about one and one-third times as long as first tergite, apex of
petiole about twice as wide as base, pygidial area with distinct lateral carinae, parallel on
apical half, gradually diverging towards base. I have seen one male, recorded above, from
Bekily (det. J. Leclercq). This differs from the male of P. reticulosus, apart from being
smaller, in having no tyloidea. Raised part of interantennal carina hardly broadened.
Arnold described the punctation of the scutum as being denser than in the female, inter-
spaces between larger punctures about twice as wide as punctures themselves and some
specimens with a trace of transverse rugosity. He did not mention the interantennal carina.

70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

The above male has a strongly transversely rugose scutum, apical margin of scutellum
yellow, remaining part brown and yellowish-brown. Petiole about one and a half times
as long as first tergite.

Psenulus reticulosus Arnold

Arnold, 1945: 154—156, 9 and & (Psenulus reticulosus and Psenulus reticulosus var. albo-
scutellatus; Malagasy: Rogez, Perinet, Ivondro, Ranomafana).

Leclercq, 1961a: 35, 2 (Malagasy: Mandraka, Rogez, Analandraraka).

Leclercq, 1961c: 117, 9 (Malagasy: Ranomafana, Perinet, Joffre-ville).

Material studied: Rogez, 1 9, Jan., 1931, 1 2,1932, 1 S, April, 1931 (FAG); 1 9,
Rogez, “forêt côte est”, Febr., 1934, coll. A. Seyrig (author's collection); Rogez, 1 9,
Dec., 1931, 1 9, 1932, 1 g', May, 1932 (FAG). The last mentioned three specimens
belong to the var. alboscutellatus Arnold.

Petiole over one and a half times as long as first tergite, in dorsal aspect. Prescutal
sutures distinct only on basal third of scutum. Raised part of interantennal carina slightly
broadened in female, hardly or not broadened in male.

Pygidial area of female narrow with indistinct, parallel, carinae.

Underside of fore and mid femora pale brown in male, in female fore and mid legs
completely pale yellow. Antennal segments 3 (or 4)—13 with distinct tyloidea, which
are reddish like flagellum.

Psenulus sapobaensis spec. nov.

Female. — Head and thorax black; mandibles yellowish with brown tips, labrum
reddish-brown, palpi testaceous. Scape of antennae yellowish, following segments dark
brown above, last segment reddish-brown above, underside of antennae yellowish-red.
Pronotal dorsum medially, and metanotum, blackish-brown. Posterior part of pronotal
tubercles yellowish, tegulae yellowish-red. Fore and mid legs including trochanters and
mid coxae pale yellowish-red or straw-yellow; hind legs including coxae darker yellowish-
red. Veins of wings dark brown. Gaster including petiole yellowish-red.

Frontal carina distinct, raised part between antennae very slightly broadened (Fig. 43),
ending below antennae in a distinct transverse carina. Supraclypeal area and basal half
of clypeus superficially densely punctate, apical half of clypeal disk not very shining but
without distinct sculpture or punctation. Protruding median part of clypeal margin
distinctly bidentate, distance between apices of teeth about one-ninth of total distance
there between the eyes (Fig. 42). Frons below anterior ocellus with distinct separate
punctures, near oculi almost smooth. Vertex shining, sparsely and minutely punctate.
Tempora with fine longitudinal striae. Occipital carina ending normally in hypostomal
carina. Mandibles normal. Third antennal segment about two and one-third times, fourth
segment about twice, fifth segment about one and three-quarter times, segments 6—8
about one and a half times, segments 9—11 about one and one-third times as long as
broad at apex, last segment at least twice as long as broad at base (all in frontal view).

Pronotal corners sharp. Scutum fairly strongly punctate, finer on anterior part between
prescutal sutures. Prescutal sutures distinct on anterior two-fifths of scutum, continued
until near apical margin by a row of relatively large punctures. Scutellum and metanotum
scarcely finely punctate. Propodeal enclosure triangular, well-defined, median part rather

J. P. VAN LITH: Revision of Ethiopian Psenini 71

47
46

Fig. 42—44. Psenulus sapobaensis sp. nov., 2, holotype. 42, face; 43, frontal carina, dorsal aspect;
44, propodeal enclosure. Fig. 45—47. Psenulus leoninus sp. nov., 9, holotype. 45, face; 46, frontal
carina, dorsal aspect; 47, pygidial area

wide with median longitudinal carina (Fig. 44), lateral parts with few oblique carinae.
Upper part of propodeum with narrow smooth and shining area, back with fine oblique
striae, transverse on lower part. Dorso-lateral edge of propodeum more coarsely reticulate,
posterior part of sides finely reticulate, anterior part smooth. Metapleura shining, some-
what striate below, mesosternum and mesopleura smooth, with scattered punctures.
Anterior oblique suture foveolate, widened upper part smooth. Legs stout, fore and mid
femora and tibiae rather heavily built, apex of mid tibiae on outer side with narrow
longitudinal depression, margined anteriorly by a reddish oblique low carina and behind
by four short reddish spines. Hind tibiae on outer side with three (left tibia) or two
(right tibia) longer spines. Second submarginal cell above about half as wide as lower
side. First recurrent vein ending just in second submarginal cell, second recurrent vein
ending well in third submarginal cell, about one-fifth of base of third submarginal cell.
Petiole long, slightly more than one and a half times length of first tergite, cylindrical,
apex dorsally with small triangular pit. Gaster minutely punctate, no well-defined pygidial
area, just a narrow smooth and shining area bordered by a few long hairs. Base of second
sternite somewhat depressed.

Pubescence of face and tempora silvery, appressed, leaving structure of clypeus well
visible, face also with a number of long erect hairs. Vertex with long greyish pubescence.

72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

Pubescence of thorax whitish, long on propodeum, dense on mesosternum. Epicnemial
areas below with round patch of dense appressed silvery pubescence. Legs and gaster with
yellowish-grey pubescence, denser and rather long on apical margins of fourth and fifth
sternites but not fringe-like, sixth sternite densely yellowish-golden pubescent. Petiole with
a few long, obliquely directed downwards, hairs, dorsum bare.

Length about 9 mm.

Male unknown.

Nigeria: 1 ® (holotype), Sapoba, 16 Aug., 1962, coll. D. C. Eidt, in Malaise trap
(CNC).

This species is probably closely related to P. reticulosus.

Psenulus leoninus spec. nov.

Female. — Head and thorax black; mid part of mandibles brownish, palpi testaceous,
antennae dark-brown, scape and underside of following segments reddish-brown. Pronotal
tubercles slightly brownish, tegulae reddish. Fore and mid legs reddish, including tro-
chanters, mid femora brown below, fore and mid basitarsi yellowish; hind trochanters
brown, hind femora brown, apical two-thirds reddish below, rest of hind legs also
reddish. Veins of wings dark-brown. Petiole including ventral plate black; gaster reddish,
first tergite with large black mark almost reaching hind margin.

Frontal carina raised and widened between antennae, about two and a half times as
long as broad (Fig. 46), ending below antennae in a short transverse carina, in the shape
of a reversed V with obtuse angle. Face below transverse carina dull, finely aciculate;
clypeal margin ending in two distinct small teeth, distance between these teeth about one
eighth of total distance there between eyes (Fig. 45). Mandibles normal. Occipital carina
ending in hypostomal carina. Frons densely punctate, vertex between eyes with more
widespread punctures. Tempora aciculate, partly finely striate. Antennae short, third
segment about one and a half times, segments 4—6 about one time, segments 7—9 about
two-third times, segments 10—11 about half as long as broad at apex; last segment about
one and a third times as long as broad at base.

Pronotal lateral corners with sharp anterior angle. Scutum finely densely punctate,
relatively few very large punctures and on sides of anterior half transversely striate.
Prescutal sutures almost reaching hind margin of scutum, rather ill-defined. Scutellum
and metanotum with sparse fine punctures. Propodeal enclosure narrow, lateral parts with
a few oblique carinae. Median longitudinal groove wide, with transverse carinae. Upper
part of back of propodeum immediately behind enclosed area finely obliquely striate,
lower half and posterior two-thirds of sides of propodeum with irregular reticulate carina-
tion; anterior part of sides dull. Metapleura shining. Mesopleura and mesosternum with
few fine punctures. Anterior oblique suture coarsely foveolate, widened upper part with
a few transverse carinae. Legs normal, flattened apical outer side of mid tibiae with short
row of three small reddish teeth. First recurrent vein of fore wings interstitial, upper side
of second submarginal cell about half length of lower side. Petiole little longer than first
tergite, cylindrical, apex dorsally with long triangular emargination and small pit, anterior
third dorsally flattened and medially with slight groove. Gaster very finely punctate,
pygidial area narrow-triangular, lateral carinae parallel on apical half (Fig. 47).

Face below antennae with appressed silvery pubescence and erect silvery hairs. Vertex
with long yellowish-grey pubescence, not dense, rest of head and thorax with whitish

J. P. VAN LITH: Revision of Ethiopian Psenini 73

pubescence, pubescence on mesosternum dense and short, also with long hairs; hairs on
scutellum, metanotum and propodeum long. Epicnemial areas below with round patch of
dense yellowish-white pubescence. Petiole with a few long laterally outstanding hairs.
Gaster yellowish-grey pubescent, hind margins of fourth and fifth sternites with dense
fringe of yellowish-golden hairs, at least as long as width of base of hind basitarsus.
Sixth sternite densely yellowish-golden pubescent.

Length about 6 mm.

Sierra Leone: 1 9 (holotype), Freetown, August, 1967, coll. D. F. Owen (HT).

P. leoninus may be closely allied to P. /usingae Leclercq, described from Zaire, but it is
easily distinguished by the smaller size, the shape of the lower part of the frontal carina,
the shorter petiole, the coarser sculpture of the scutum, the paler legs and the darkened
first gastral tergite.

Psenulus benoiti Leclercq
Leclercq, 1961a: 29—30, 9 and & (Zaire).

The female of P. benoiti has a remarkable sharp median longitudinal carina on the
sixth tergite, as if the lateral carinae of the pygidial area have joined into one single
carina. This characteristic is not found in any of the other Old World Psenini, only in
the female of the Philippine P. /uteopictus the sixth tergite is very slightly keeled. P.
mayorum Bohart & Grissell (1969) from Mexico, shows a similar structure of the sixth
tergite, but it certainly belongs to a different group of species.

A more extensive description of P. benotti is given here.

Female. — Head and thorax black; antennae yellowish-brown below, brown above,
mandibles reddish-brown, palpi testaceous. Pronotal tubercles and tegulae yellowish-red.
Fore and mid legs including trochanters yellowish, back of femora brownish. Hind
trochanters yellowish, femora brown, base of hind tibiae and tarsi yellowish-brown, apex
of hind tibiae brownish. Petiole black except ventral plate, gastral tergites 1—6 reddish.
Veins of wings brown.

Frontal carina between antennae hardly broadened, but with distinct narrow groove.
Distinct transverse carina below antennae. Narrow protruding median part of clypeal
margin not distinctly emarginate. Frons shining, finely punctate, vertex more shining,
almost impunctate. Tempora finely alutaceous, almost smooth. Occipital carina ending in
hypostomal carina. Third antennal segment about twice as long as broad at apex, follow-
ing segments gradually shorter, segments 9—11 much broader than long, twelfth segment
about one and three-quarters times as long as wide at base. Mandibles normal.

Punctation of scutum variable, on median part of scutum large impunctate areas.
Prescutal sutures almost reaching hind margin, on fore part with some transverse striae,
parapsidal sutures long. Enclosed area of propodeum short, deep, median area broad but
much less than in the group of P. turneri. Longitudinal propodeal suture widened on
back-side. Back of propodeum smooth and shining, apex with some irregular carination;
sides of propodeum on posterior part coarsely reticulate, almost smooth on fore part.
Metapleura shining; anterior plate of mesepisternum, mesopleura and mesosternum
sparsely finely punctate. Anterior oblique suture foveolate, widened upper part indistinct.
Legs normal. Second submarginal cell above about two-thirds of length of lower side.
Petiole long, in dorsal view about one and a half times as long as first tergite, cylindrical,

74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

base dorsally somewhat flattened, apex with deep triangular pit. Gaster finely punctate.
No normal pygidial area, but sixth tergite with sharp narrow median carina, which ends
in an indistinct very small elongate and flattened part. Second gastral sternite with semi-
circular depression.

Pubescence of face silvery, appressed, leaving sculpture visible. Head and thorax
whitish pubescent, dense on mesosternum, petiole with long, laterally and ventrally out-
standing yellowish-grey hairs, gaster with yellowish pubescence. Epicnemial areas below
with circular densely whitish pubescent spot, last sternite with dense yellowish-golden
pubescence, apical margins of fourth and fifth sternites with short dense pubescence, not
conspicuous.

Length about 6.5 mm.

Male. — Resembling female. Fore and mid legs including trochanters more reddish,
hind trochanters, femora, and tibiae dorsally, brown.

Frons on both sides of median carina somewhat convex. Frons with fine distinct
punctation. Frontal carina between antennae raised and broadened, ratio width-length
about 1 : 3. Antennal segments 3—8 about one and a half times, segments 9—12 about
one and one-third times as long as broad at apex, last segment about twice as long as
broad at base. No distinct tyloidea, lower side of antennal segments distinctly convex.
Median part of clypeal margin with two distinct small teeth. Prescutal sutures without
distinct transverse striae, almost reaching hind margin and ending in a widened depres-
sion. Upper part of propodeum shining with a few indistinct punctures, back coarsely
reticulate, posterior part of sides of propodeum also coarsely reticulate, fore part densely
punctate. Petiole about one and one-third times as long as first gastral tergite, sides with
lower carina on apical half and distinct longitudinal depression, dorsal side rounded,
flattened at base and with triangular pit at apex.

Length about 6.5 mm.

Material studied: © (holotype) and & (allotype), Zaire, Upper Uele, Isiro (Paulis),
May and July, 1947, coll. P.L.G. Benoit (MRAC).

No further material seems to have been collected thus far.

Psenulus kohli Arnold

Arnold, 1923: 50—51, 9 (Rhodesia: Bulawayo).
Leclercq, 1961a: 30.

After study of the holotype (RM) a few remarks should be made. — Femora brown,
tibiae and tarsi whitish, almost whole underside of fore tibiae light brown, mid tibiae
light brown below except apices, apical half of hind tibiae brown, except a narrow
yellowish-white line on outer side. Greater part of mandibles reddish.

Raised part of carina between antennae much broadened, with distinct lateral carinae,
broadened part closed below by indistinct converging carinae on lower part. This does
not appear from Arnold’s figure (1923, Fig. 77, p. 51). Tempora broad. Upper side of
second submarginal cell about two-fifths of length of lower side of cell. Propodeal
enclosure very short, behind this area very fine oblique striation. Petiole cylindrical, at
least one and a half times as long as first tergite, in dorsal aspect. Pygidial area very
narrow elongate-triangular, lateral carinae distinct.

J. P. vAN LITH: Revision of Ethiopian Psenini 75

Also broadened part of interantennal carina silvery pubescent. Apical margins of fourth
and fifth sternites with short pubescence.

This species is not closely allied to P. capensis and P. latiannulatus, as Arnold sug-
gested, but it takes an isolated position. The gaster is not slender and the first tergite is
hardly nodose. P. kohli is easily distinguished from other Ethiopian species by the absence
of a transverse carina below the antennae and the shape of the densely silvery pubescent
interantennal carina.

Psenulus luctuosus Arnold

Arnold, 1929: 405—-406, 9 (Rhodesia).
Leclercq, 1961a: 30.

Supplementary description of holotype: 9, Vumba Mts., Umtali, Rhodesia, Febr.,
1928, coll. G. Arnold (RM). — Pronotal tubercles dark brown. Interantennal carina not
quite closed below, as Arnold’s figure No. 37 (1929, p. 406) suggests, but slightly open
and the two lateral carinae ending close together in transverse carina (cf. Fig. 48: P.
jacoti). Anterior oblique suture strongly foveolate, widened upper part with transverse
carinae. Petiole more than twice as long as first gastral tergite. Pygidial area elongate-
triangular, lateral carinae slightly diverging towards base, surface irregularly finely
punctate.

Petiole with long erect lateral hairs.

No further captures have come to my knowledge.

This species is closely allied to P. jacoti sp. nov., from which it differs in the reddish
fifth and sixth tergites, the longer petiole and the slightly wider pygidial area.

Psenulus stevensoni Arnold

Arnold, 1940: 133—134, Q and & (Rhodesia: Bulawayo).
Leclercq, 1961a: 30.

Material studied: 1 ©, Rhodesia, Bulawayo, Hillside, 29 April, 1927 (with red type-
label); 1 g', Bulawayo, 9 Nov., 1924 (with red type-label); 1 g', Bulawayo, Hillside,
29 April, 1927, all coll. R. H. R. Stevenson (RM). I have selected the female as the
lectotype and the male of 9 Nov., 1924, as the paralectotype. Arnold apparently has not
seen the male which was collected at the same time and locality as the female.

Supplementary description of female. — Not only basitarsi of mid legs yellowish-white,
but also basitarsi of fore legs more or less yellowish. Hind legs more brown than
brownish-red. Scape of antennae, following four segments completely and segments 6—9
below reddish, rest of flagellum dark brown.

Broadened part of interantennal carina open below, lateral carinae ending in transverse
carina (see Fig. 48: P. jacoti, and Leclercq, 1961a). Scutum dull, superficially transversely
striate and also with some rather large punctures. Petiole about two and a half times as
long as first tergite. Pygidial area distinct, elongate-triangular.

Apical margins of fourth and fifth sternites with fringe of short white hairs.

Supplementary description of male. — Basitarsi of fore and mid legs reddish.
Broadened part of interantennal carina narrower than in female, closed below and ending
as a narrow single carina in transverse carina below antennae. Antennal segments 3—13
with narrow oblique tyloidea. Scutum more strongly transversely rugose. Parapsidal

76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

sutures deep, as in female. Petiole more than twice as long as first tergite, in dorsal aspect.

For further characteristics may be referred to Arnold (1940).

Arnold considered P. stevensoni closely allied to P. kohli, but in differs from the latter
species in having a distinct transverse carina below the antennae in the female, the head
is much swollen behind the eyes, the propodeal enclosure is longer, and the petiole is
much longer. There are much closer relationships with P. lwctwosus Arnold, P. jacoti
sp. nov., and perhaps also with P. /usingae Leclercq and P. rugifrons sp. nov.

On the label of the female its prey is glued, probably an adult Psyllid which is some-
what damaged.

Psenulus lusingae Leclercq
Leclercq, 1961a: 33, 2 (Zaire).

The characteristics given by Leclercq in his key (1961a) are sufficient to recognize this
species. However, a redescription including some not yet mentioned details may be useful
for future study.

Female. — Head and thorax black; mandibles dark reddish except tips, palpi testaceous,
scapes of antennae and underside of flagellum reddish-brown, dorsally almost black.
Tegulae reddish-brown. Veins of wings brown. Legs brown, fore side of fore femora,
tibiae and whole tarsi yellowish-red, mid basitarsi and second tarsal segment testaceous.
Petiole including ventral plate black, rest of gaster reddish.

Raised part of frontal carina between antennae broadened, ratio width-length about
1 :3, lateral carinae of broadened part ending separately, parallel, in transverse carina
below antennae, this transverse carina almost rectangular in frontal view. Frons below
ocelli finely punctate, vertex nearly impunctate. Mandibles normal. Protruding median
part of clypeal margin with two distinct teeth, distance between apices about one-seventh
of total distance there between eyes. Tempora very finely longitudinally striate. Antennae
short, somewhat clavate, third segment about one and three-quarters times, segments 4—5
about one and one-third times, segments 6—8 about as long as wide at apex, segments
9—10 about two-thirds times, eleventh segment about half as long as wide at apex, last
segment about one and a half times as long as wide at base.

Scutum opaque, very finely punctate, interspaces a few times larger than punctures.
Prescutal sutures nearly reaching hind margin of scutum, not well-defined, parapsidal
sutures much deeper. Scutellum and metanotum shining, sparsely punctate. Propodeal
enclosure triangular, central area normal, lateral areas each with about six oblique carinae,
propodeum behind enclosed area very finely and densely obliquely striate, median longi-
tudinal sulcus gradually narrowing towards centre, followed by a few narrower alveoles.
Also greater part of back-side of propodeum with very fine oblique striae, dorsolaterally
a coarsely reticulate-carinate ridge, sides of propodeum on posterior half with fine
reticulation, on fore part almost smooth, with fine punctures. Metapleura shining,
mesopleura and anterior plate of mesepisternum extremely finely punctate. Anterior
oblique suture coarsely alveolate, widened upper part with a few transverse carinae. Upper
side of second submarginal cell about one-quarter of length of base. Legs normal, outer
apex of mid tibiae flattened, this area defined posteriorly by about six small thorns.
Petiole slender, cylindrical, over twice as long as first tergite, in dorsal aspect, dorsally
rounded with small pit at apex. Pygidial area narrow, distinct. Second gastral sternite
with indistinct basal depression.

J. P. VAN LITH: Revision of Ethiopian Psenint 77

Face with appressed silvery pubescence, head, thorax and legs whitish pubescent, back
of propodeum below with longer hairs. Petiole laterally with long erect hairs. Gaster with
short yellowish-grey pubescence, apical margins of fourth and fifth sternites with dense
and short pale pubescence, sixth sternite densely yellowish-golden pubescent.

Length about 7.75 mm.

Material studied: 9 (holotype), Zaire, Parc National de l’Upemba, Lusinga, 1.760 m,
3 July, 1947, coll. Mission G. F. de Witte, no. 542a (MRAC).

No further specimens are known at present.

Psenulus avernus Leclercq

Leclercq, 1961a: 30, 4 (Zaire).

Supplementary description of holotype, from Rutshuru (IRSNB). — Frons shining,
finely densely punctate, vertex almost impunctate. Tempora finely striate below. Antennal
segments 3—11 with distinct narrow tyloidea, small and indistinct on twelfth segment.

Prescutal sutures long, almost reaching hind margin of scutum. Upper part of pro-
podeum shining behind enclosed area, with irregular oblique striation. Back-side of
propodeum with coarse reticulate carination. First recurrent vein ending just in second
submarginal cell, second recurrent vein well in third submarginal cell, this cell above
about half as wide as lower side of cell. Petiole long and slender, about two and one-
third times as long as first tergite in dorsal aspect, laterally somewhat depressed, on
apical two-thirds with indistinct upper and distinct lower keel.

Estimated length — the gaster has been glued to the pin — 7 mm.

P. avernus is probably closely related to P. /usingae Leclercq, of which only the female
holotype, from Lusingae, Zaire, is known. However, this female has a very densely and
finely striate upper part of the propodeum; its scutum is rather dull, this part being
shining in P. avernus.

Psenulus jacoti spec. nov.

Female. — Head and thorax black; following parts reddish-brown: Mandibles except
dark tips, antennae including scape, but segments 7—11 dorsally somewhat darkened,
pronotal tubercles and tegulae. Palpi brown. Fore legs reddish-brown, greater part of
femora below and back of trochanters black; mid trochanters and basal half of mid
femora black, rest of mid legs reddish-brown; hind trochanters and femora except apex
ventrally, black, hind tibiae and tarsi reddish-brown, partly blackish. Veins of wings
almost black, upper half of radial cell infuscate. Gaster black, petiole ventrally and later-
ally, at least partly, reddish-brown, also first tergite somewhat on sides of apical part,
apex of last segment reddish-brown or yellowish-red.

Frontal carina raised and much broadened between antennae (Fig. 49), narrowed below
antennae into two carinae which are closely parallel or just touching, and ending in a very
fine, almost indistinct, angular transverse carina (Fig. 48). Protruding median part of
clypeus with two distinct triangular teeth, distance between apices of teeth about one
seventh or one eighth of total distance there between eyes. Frons shining, finely and
closely punctate, between oculi and ocelli much sparser punctate, behind ocelli almost
impunctate. Tempora, especially lower part, with dense fine oblique striation. Apex of
mandibles normally bidentate. Antennae widening towards apex, third segment about

78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

Fig. 48—51. Psenulus jacoti sp. nov., ®, paratype. 48, face; 49, frontal carina, dorsal aspect; 50,

2nd and 3rd submarginal cell of right fore wing; 51, pygidial area. Fig. 52—54. Psenulus jacoti

sp. nov., 6, paratype. 52, 7th sternite, lateral aspect; 53, 8th sternite, lateral aspect; 54, genitalia,

dorsal jaan Fig. 55—58. Psenulus rugifrons sp. nov. ®, holotype. 55, face; 56, frontal carina,
dorsal aspect; 57, 2nd and 3rd submarginal cell of right fore wing; 58, pygidial area

J. P. van LITH: Revision of Ethiopian Psenini 79

twice, segments 4—5 about one and a half times, segments 6—7 about one and a fourth
times as long as broad at apex, segments 8—11 nearly quadrate, last segment about one
and three-quarters times as long as broad at base.

Lateral corners of pronotum obtuse. Scutum shining, sparsely punctate, prescutal
sutures long, on posterior margin lost in the denser punctation. Scutellum and meta-
notum almost impunctate. Enclosed area of propodeum rather short, shining, with oblique
carinae. Dorsal half of back of propodeum finely and densely obliquely striate, striae
reaching enclosed area, lower part of propodeum with fine sculpture and punctures.
Median longitudinal carina broad above, narrowing towards apex, upper part with
transverse carinae. Sides of propodeum with fine oblique striae. Metapleura and meso-
pleura shining, sparsely punctate, mesosternum somewhat more densely punctate. Anterior
oblique suture foveolate, widened upper part with transverse carinae. Legs rather stoutly
built, mid tibiae flattened on outer side of apex, with a few dark red spines at posterior
margin, hind tibiae with a row of short reddish spines on outer side. Second submarginal
cell pentagonal (Fig. 50), first recurrent vein ending in second submarginal cell, second
recurrent vein ending just in third submarginal cell, in some specimens interstitial. Petiole
almost cylindrical, about one and a half times as long as first tergite (in dorsal view),
dorsally slightly flattened, with small pit at apex. Gaster very finely punctate, segments
2—6 finely aciculate. Pygidial area narrow (Fig. 51), lateral carinae distinct, almost
parallel, medially a line of fine punctures.

Face with appressed silvery pubescence and long, erect, silvery hairs; rest of body
greyish-yellow pubescent. Epicnemial areas below with round patch of dense, yellowish-
grey pubescence, appressed, apical margins of sternites 4—5 with dense, short, whitish
pubescence, sixth sternite with long and dense, directed backwards, pale golden pubes-
cence. Pubescence on mesosternum rather long and dense.

Length about 7 mm.

Male. — Similar to female but reddish-brown parts more or less darkened. In two of
the three males antennal segments 3—13 black above, in one of the two males from
Belmont pronotal tubercles and mid and hind legs almost completely black. In the allo-
type the antennae are almost completely reddish.

Transverse carina below antennae more distinct. Raised carina between antennae less
broadened. Antennal segments 3—12 about one and a half times as long as broad at
apex, last segment twice as long as broad at base; segments 3—11 with narrow longi-
tudinal tyloidea, 12th segment with small, indistinct, tyloidea. Seventh sternite with small
triangular emargination at apex (Fig. 52). Eighth sternite: Fig. 53. Genital apparatus
(Fig. 54) short, pale brown, lateral valves with inner tooth, apices with rounded outer
margin.

No conspicuous pubescence on epicnemial areas and on margins of fourth and fifth
gastral sternites.

Length about 6.5—7 mm.

South Africa, Cape Province: 1 9 (holotype), Hilton, Grahamstown, 21 Oct., 1970,
coll. C. Jacot-Guillarmod; 1 & (allotype), Howison’s Poort, Grahamstown, 19—22
Nov., 1971, coll. F. W. Gess; paratypes: 3 9, Hilton, Grahamstown, 5—9 Nov., 1970,
coll. F. W. Gess, 1 9, Hilton, Grahamstown, 19—31 Dec., 1970, C. Jacot-Guillarmod,
1 9, Howison’s Poort, Grahamstown, 12—14 Nov., 1971, F. W. Gess, 1 © , Kenton-on-
Sea, Dec. 1971, R. A. Jubb, 2 g', Belmont Valley, Grahamstown, 28—31 Oct., 1971,
coll. F. W. Gess, all in Malaise traps (AMG, one $ and one g' in author's collection).

80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

South Africa, Transkei: 1 9, Port St. John, 12 Nov., 1961, coll. H. N. Empey (EC).

There is some variation in this species. In one of the females from Hilton the frontal
carina is not bicarinate below the antennae, but single. In the female from Kenton-on-
Sea the petiole is abnormally broadened towards the apex. The female from Transkei is
paler coloured: antennae dorsally hardly darkened; fore and mid femora and trochanters
brown instead of black; hind trochanters dark-brown, femora brown, hind tibiae and tarsi
reddish-brown; complete petiole and great part of first tergite dark reddish; veins of
wings brownish.

P. jacoti seems to be closely related to P. stevensoni Arnold from which it can be
distinguished, e.g. by the non-striate vertex. There is also a great similarity to P. lwctuosus
Arnold, P. /usingae Leclercq, and P. rugifrons Van Lith, which all have at least the fifth
and sixth tergites reddish.

Psenulus rugifrons spec. nov.

Female. — Head and thorax deep black; apices of mandibles dark reddish, labrum and
palpi dark brown, underside of antennal segments 5—12 with orange-brown spot. Legs
black; tarsi, underside of fore tibiae and base of hind tibiae brown, apical spine of fore
tibiae yellowish-brown, of mid legs whitish, inner apical spine of hind legs whitish,
outer spine of hind tibiae brown. Petiole black, extreme apex dorsally, ventral plate
completely and whole gaster reddish.

Frontal carina raised and broadened between antennae (Fig. 56), broadened area
tapering to two parallel carinae which end in a transverse carina, reversed V-shaped
(Fig. 55). Face below antennae, and clypeus, dull, finely punctate, median part of
clypeus protruding, with two small teeth close together. Frons finely obliquely punctato-
rugose, between eyes and ocelli finely and sparsely punctate, vertex finely rugose.
Tempora finely striate. Third antennal segment about twice, segments 4—5 about one
and one-third, sixth segment about one and one-fourth times as long as broad at apex,
segments 7—9 about as long as broad, segments 10—11 broader than long, last segment
at least one and a half times as long as broad at base. Mandibles normal.

Anterior lateral corners of pronotum obtuse. Scutum finely punctate, prescutal sutures
almost reaching hind margin, not distinctly alveolate, parapsidal sutures distinct, broad.
Lateral margin of scutum along tegulae strongly alveolate. Scutellum finely punctate,
metanotum almost impunctate. Propodeal enclosure shining, with distinct oblique carinae
on lateral parts, median part of usual width. Upper part of propodeal longitudinal
sulcus broad, with a few transverse carinae, sulcus on back side of propodeum broad but
not deep, dull. Fine oblique striation behind propodeal enclosure, lower part finely
reticulate; propodeum postero-laterally coarsely reticulate, sides of propodeum posteriorly
finely reticulate, anteriorly very finely striate. Metapleura somewhat shining, mesopleura
and mesosternum shining, finely punctate. Anterior oblique suture foveolate, upper part
with transverse striae. Legs normal. First recurrent vein of fore wings ending well in
second submarginal cell, second recurrent vein in third submarginal cell; upper side of
second submarginal cell about one-fourth of length of lower side (Fig. 57). Petiole
cylindrical, nearly twice as long as first gastral tergite, apex dorsally with long triangular
depression. Pygidial area distinct, narrow with almost parallel carinae (Fig. 58).

Face with short and long, silvery, appressed pubescence. Head and thorax greyish-

J. P. VAN LITH: Revision of Ethiopian Psenint 81

white pubescent, on mesosternum longer and denser. Petiole dorsally along sides with
row of short hairs, laterally with long outstanding hairs. Gaster yellowish-golden pubes-
cent, longer on last segment. Apical margins of fourth and fifth gastral sternites with
dense fringe of short pale hairs, sixth sternite with long, backwards directed, yellowish-
white hairs.

Length about 7 mm.

Male unknown.

Ethiopia: 1 2 (holotype), Jimma, 29 June—3 July, 1965, coll. A. B. Gurney
(USNM).

P. rugifrons probably belongs to the group of P. J4ctuosus Arnold.

Psenulus bidentatus (Cameron)

Cameron, 1910: 282, 9 (Psen bidentatus; Tanzania: Kilimanjaro).

Turner, 1912: 412—413, 2 (Psenulus rubrocaudatus; South Africa: Natal).
Arnold, 1923: 48—50, 9 (Zaire: “Elizabethville” (Bequaert)).

Leclercq, 1961a: 33—34, 9 and & (Zaire).

Leclercq, 1961b: 58, 9 and & (Zaire).

P. bidentatus is recognized by the bidentate clypeus, slightly broadened raised part of
interantennal carina, distinct transverse carina below antennae, finely striate tempora and
long petiole. The pygidial area of the female is indistinct, the male has very strongly
moniliform antennae and the raised part of the interantennal carina is narrow in most
cases.

A study of the holotype (female) from the Kilimanjaro, which is in the Natural
History Museum at Stockholm, revealed that most of the specimens from Zaire (recorded
as P. bidentatus Cameron by Leclercq), part of the material from Rhodesia, and specimens
from Malawi and Equatorial Guinea belong to a different form. There are distinct
differences in structure as well as in colour and size and I believe that they represent at
least a distinct subspecies. Future studies may lead to the conclusion that pallidus has
specific rank.

Leclercq (1961a) pointed out that P. rubrocaudatus Turner is a geographical form of
P. bidentatus in South Africa.

Psenulus bidentatus bidentatus (Cameron)

Cameron, 1910: 282, ® (Psen bidentatus; Tanzania: Kilimanjaro).
Arnold, 1923: 48—50, 9 (Psenulus bidentatus; Rhodesia: Salisbury).
Arnold, 1929: 406 (Fig. 38: head of 9).

Leclercq, 1961a: 33—34, 9 partim (Zaire).

Most of the specimens recorded by Leclercq from Zaire (1961a, b) belong to the
subspecies pallidus. The following female from Lubumbashi (Elisabethville) is a repre-
sentative of the nominate form: Lubumbashi, Oct., 1934, coll. P. Quarré (MRAC).

New records: Rhodesia: 4 2, Salisbury, Sept., 1916, coll. G. Arnold, 26 Sept., 1919,
14 May, 1941 (2 specimens); 2 9, Bulawayo “Kh” (Khami?), 21 Oct. 1938, coll.
R. H. R. Stevenson (RM).

Kenya: 2 g', Nairobi, July and Aug., 1930, coll. V. G. L. van Someren (BM, 1972-2,
pres. by Com. Inst. Ent., and BM 1959-468).

82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

Angola: 1 © and 1 g', 30 km N of Quiculungo, Sept—Oct., 1957, George R. Fer-
guson Collection 1972 (OSUM).

Probably the first-mentioned female from Salisbury, labelled “Psenulus Bequaerti
Brauns comp. with type, G.A.”, was the one recorded by Arnold (1923). He compared
it with the type of P. beguaerti Brauns (nomen in litt.) from Lubumbashi and stated that
they differ only in minor details of colour. I presume Brauns’ specimen was a female of
pallidus subsp. nov.

The authorities of the Naturhistoriska Riksmuseet at Stockholm kindly allowed me to
study the holotype (female), labelled “Kilimandj., Sjöstedt, Kibonoto Kulturz., 4 maj”
(printed) and “Psen bidentatus Cam. type” (in Cameron’s handwriting).

Supplementary description of female. — Labrum reddish. Fore and mid trochanters
brown, fore and mid legs reddish but lower side of femora brown, sometimes foreside of
fore tibiae and extreme base of mid tibiae yellowish, outer side of mid tibiae pale brown
and basal two-thirds of mid basitarsi, sometimes also fore basitarsi, yellowish. Hind legs
brown, femora almost black. Tegulae reddish, very rarely somewhat yellowish. First
gastral tergite except apical margin and base of second gastral tergite dark brown; in the
female from Angola also first tergite reddish.

Raised part of interantennal carina slightly broadened, lateral ridges of broadened part
shining, interantennal carina ending below antennae in a distinct transverse carina. Frons
densely finely punctate, almost smooth near eyes. Vertex finely punctate. Apical third of
clypeus sparsely punctate. Tempora very finely striate on lower part. Third antennal seg-
ment about two and a half times, fourth segment about one and three-quarters times, fifth
segment about one and a half times as long as broad at apex, segments 6—7 about as
long as wide, segments 8—11 shorter than wide, last segment about one and a half times
as long as broad at base.

Pronotal corners obtuse. Punctures of anterior plate of mesepisternum somewhat
elongate, mesopleura strongly punctato-striate, apex below finer and more sparsely
punctate. Punctation of scutum finer, interstices mostly larger than punctures, very light
tendency to transverse striation. Prescutal sutures almost reaching hind margin, fore part
more linear, posterior half more foveolate. Anterior oblique suture foveolate, widened
upper part with a few transverse carinae. Oblique carinae of propodeal enclosure weak.
Propodeum behind enclosed area with narrow smooth margin, upper part of back of
propodeum obliquely, lower part more irregularly striate. Petiole about twice as long as
first tergite. Pygidial area not sharply defined, dull, indicated by two almost parallel low
ridges, apex blunt, as in P. nigeriae Leclercq.

Length about 8.5—9.5 mm.

First description of male of nominate form. — Resembling female in colour and
structure. Fore and mid basitarsi reddish or yellowish-red. First tergite except apical
margin dark brown. Scape and following segments of antennae reddish, flagellum dark
brown above, apical half of last segment above reddish-brown, underside of flagellum
reddish.

Raised part of interantennal carina very slightly or not broadened. Antennae strongly
moniliform, “en forme de toupies’ (Leclercq, 1961a), segments 3—13 with narrow
oblique tyloidea on back-side, segments 3—9 about one and one quarter times as long
as greatest width, segments 10—11 about as long as greatest width, last segment over
twice as long as greatest width. Scutum slightly more coarsely sculptured. Propodeum

J. P. VAN LITH: Revision of Ethiopian Psenini 83

behind enclosed area with fine superficial oblique striae which reach enclosed area; in the
male from Angola a very narrow smooth margin behind enclosure.
Length about 7.75—8 mm.

Psenulus bidentatus (Cameron) var. or subsp. ?

Ethiopia: 1 9, Adola, “S. Abyssinia”, Nov., 1941, 7000 ft, “Psenulus bidentatus Cam.
var. n., det. G. Arnold” (RM).

In this dark form of P. bidentatus (Cameron) the second and third gastral tergites are
also largely blackish and the legs are slightly darker reddish-brown than in the nominate
subspecies. More material is needed to decide whether it belongs to a different subspecies
or is merely a variety.

Psenulus bidentatus pallidus subsp. nov.

Leclercq, 1961a: 33—34, Q and & partim (Psenulus bidentatus; Zaire).
Leclercq, 1961b: 58, 2 and 4 (Psenulus bidentatus; Zaire).

Female. — Mandibles yellowish. Antennae brown above, reddish-brown below. All
trochanters brown. Fore femora brown, apex and foreside yellowish-red, sometimes apex
pale yellow, fore tibiae yellowish, brown below, fore basitarsi yellowish-white, rest of
fore tarsi yellowish-brown. Mid femora brown, apex yellowish, base and foreside of
tibiae yellowish, back-side brown, mid basitarsi yellowish-white, following segments
reddish-brown. Hind legs dark brown, base of femora below and base of tibiae paler.
Petiole black, apex of ventral plate reddish. First gastral tergite brown except a reddish
margin in the females from Lubumbashi (Elisabethville), in females from Rhodesia and
Malawi all tergites reddish. Pronotal tubercles yellowish, or yellowish-red, at least on
posterior half. Metanotum in one of the females from Malawi partly brownish.

Scutum coarsely striato-punctate, with irregular transverse rugae. Oblique striation on
upper part of propodeum medially reaching propodeal enclosure, laterally a large smooth
area.

Length about 7.5 mm.

Other characters as in nominate subspecies.

Male. — Mandibles yellow. Scape yellowish, following segments brown above,
brownish-yellow below. Fore and mid trochanters yellowish-brown, hind trochanters
brown. Fore legs yellowish, base, apex and foreside of tibiae and tarsi whitish, fore
femora slightly darkened below in two of the three specimens studied. Mid femora
brown, apex below more yellowish, mid tibiae and tarsi yellowish, foreside of tibiae and
basitarsi whitish. Hind legs brown, base of tibiae paler brown.

Pronotum mostly with two yellowish-brown or brownish marks, metanotum yellowish-
brown, reddish or brownish, rarely black, scutellum often somewhat brownish on poste-
rior margin. Petiole dorsally blackish-brown, base somewhat paler, apex of ventral plate
reddish. Tergites 1—6 or tergites 2—6 and apical margin of first tergite reddish.

Sculpture of scutum as in female, but interantennal carina hardly broadened, in most
of the males even narrow. Behind enclosed area of propodeum a rather broad smooth
margin, back-side of propodeum with irregular reticulate carination, coarser on lower
half. Mesopleura more finely and sparsely punctate than in female, interstices larger than
punctures, punctation denser below, in the male from Nkolentangan mesopleura on lower

84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

half also somewhat striate. Hypo-epimeral area shining, almost impunctate.

Length about 6.5 mm.

The male differs from the female in the yellowish or brownish marked pronotum and
metanotum (rarely dark) as well as in the distinct smooth margin behind the propodeal
enclosure.

Zaire (Congo): 1 9 (holotype), Lubumbashi, 10 Nov., 1923, coll. Ch. Seydel
(MRAC). Paratypes: 1 9, Lubumbashi, 1932, coll. De Loose (FAG); 1 g', Lubum-
bashi, 1933, coll. De Loose, 1 &, Parc National Albert, Rwindi, 1000 m, 22—24 Nov.
1934, coll. G. F. de Witte, no. 793, 1 g', Parc National de l’Upemba, Munoi, bifurca-
tion Lupiala, 890 m, 15—24 June, 1948 (MRAC); 1 g', Mabwe, Lake Upemba, 585 m,
21—28 Aug., 1947, Miss. G. F. de Witte, no. 719a (FAG); 1 ©, Parc National de la
Garamba, I/a/3, 8 May, 1950, coll. H. de Saeger, no. 497 (MRAC); 1 &, Parc National
de la Garamba, Nagero, 10 May, 1952, coll. H. de Saeger, no. 3503 (IRSNB); 1 2,
Haut-Uele, Abimva, 1925, coll. L. Burgeon (MRAC); 1 g', Haut-Uele, Isiro (Paulis),
July, 1947, coll. P. L. G. Benoit (FAG); 1 g', Rutshuru, North Kivu, Nov., 1937, coll.
J. Ghesquière, 1 8, Yangambi, 19 June, 1948, coll. P. L. G. Benoit (MRAC).

Malawi (Nyasaland): 1 9, Zamba, 18 Nov., 1943, 1 9, Mlanje, 2000 ft, 17 Dec.,
1944 (RM).

Rhodesia: 1 9, “Halfway Hotel Vic. Falls Rd” (about 140 miles on the main road
from Bulawayo to the Victoria Falls), Sept. 1957; 2 9, Selukwe, 9 Sept. and 7 Okt.,
1942 (RM).

Equatorial Guinea (Spanish Guinea): 1 g', Nkolentangan, coll. Tessmann (ZMB).

This form, especially the male, strongly resembles P. nigeriae Leclercq, of which only
the female sex is known, and perhaps they are closely related. It differs from the nominate
subspecies not only in being more yellowish marked and in having a coarser sculptured
scutum but it is also distinctly and constantly smaller. As suggested above pallidus may
be a distinct species.

Psenulus bidentatus rubrocaudatus Turner

Turner, 1912: 412—413, 9 (Natal).
Arnold, 1923: 48—50.
Leclercq, 1961a: 33—34.

Arnold (1923) placed P. bidentatus (Cameron) and P. rubrocaudatus Turner in
synonymy, but I share Leclercq’s opinion (1961) that P. rzbrocandatus should be con-
sidered a subspecies of P. bidentatus, at least as long as there is no proof that they
are specifically different. P. rubrocandatus is known only from South Africa: Natal
(Durban) and Cape Province (Grahamstown).

It differs distinctly from P. bidentatus s.str., collected in Rhodesia and northward, in
the much darker gaster and hind femora. Turner did not mention that the raised carina
between the antennae is slightly broadened. A pygidial area is defined by two rather
weak almost parallel carinae. Petiole about twice as long as first gastral tergite. Scutum
distinctly punctate, with slight tendency to transverse striation. In one of the paratypes
the fine oblique striae on the upper part of the propodeum reach the propodeal enclosure,
on either side of the longitudinal sulcus, whilst in the fresh specimens there is a narrow
smooth margin behind the enclosed area, followed by fine oblique striae, the back-side
being coarsely reticulate.

J. P. vAN LITH: Revision of Ethiopian Psenini 85

Mandibles almost completely reddish, also labrum. Narrow apical margin of fourth
gastral tergite reddish, in the female from Durban this margin much broader. Fore and
mid femora only blackish below.

Male still unknown.

New records: South Africa, Natal: 1 ©, Durban, June, 1959 (RM); Cape Province:
3 ©, Howinson’s Poort, Grahamstown, 22—24 Nov., 24—26 Nov. and 8—14 Dec.,
1971, coll. F. W. Gess, Malaise traps (AMG).

I had the opportunity to compare these specimens with the holotype and two paratypes
(all females) from Natal (BM).

Psenulus nigeriae Leclercq
Leclercq, 1961a: 34, 9 (Sierra Leone: Njala; Nigeria).

New records from Sierra Leone: 1 9, Njala, Dec., 1935, coll. E. Hargreaves, pres.
by Com. Inst. Ent. BM 1973—1 (BM); 1 ©, Freetown, May, 1967, coll. D. F. Owen
(HT).

The yellow marking of the female from Freetown is slightly paler than in the holo-
type from Njala, 21 Aug., 1929, coll. E. Hargreaves (BM). I have also seen the paratype,
a female from S. Nigeria, Oshogbo, Dr. T. F. G. Mayer (BM).

Supplementary description. — Antennae dorsally dark brown, reddish below, scape
pale yellow. Pronotum with two vague yellowish-brown marks, pronotal tubercles and
metanotum, except narrow lateral parts, pale yellow, fore and mid legs including
trochanters yellowish-white with vague brown streak on back; hind legs brown,
trochanters, underside of femora and base of tibiae paler. Petiole black, ventral plate of
petiole and gaster reddish.

Median part of clypeal margin distinctly bidentate, distance between apices of teeth
about one-sixth or one-seventh of total distance there between the eyes. Frons very finely
punctate. Raised part of frontal carina between antennae with small broadened part,
medially depressed (Fig. 60). Distinct transverse carina below antennae (Fig. 59).
Tempora finely aciculate. Third antennal segment about two and a half times, fourth
about twice, segments 5—6 about one and three-quarter times, seventh segment about
one and a half times as long as broad at apex, segments 8—9 about as wide as long,
twelfth segment slightly more than twice as long as broad at base.

Pronotal corners sharp (Fig. 61). Scutum with distinct punctures of variable size and
very slight tendency to transverse striation. Prescutal sutures long, almost reaching hind
margin, posterior two-thirds consisting of a row of larger punctures. Behind propodeal
enclosure a narrow smooth area, upper part of back-side of propodeum finely obliquely
striate, lower part rather finely irregularly reticulate. Apex of mid tibiae on outer side
with small bare area, raised and shining in the middle and margined posteriorly and at
apex by a few short reddish spines. Petiole nearly twice as long as first tergite, in dorsal
aspect. Pygidial area distinct, almost rectangular, apex blunt (Fig. 62), parallel carinae
not sharp, surface dull and medially with two parallel rows of very fine punctures, each
with a long erect fine hair.

Apical margins of fifth and sixth sternites with dense short pale pubescence.

Length about 8 mm.

Male unknown.

86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

Fig. 59—62. Psenulus nigeriae Leclercq, 9, Sierra Leone. 59, face; 60, frontal carina, dorsal aspect;

61, pronotum; 62, pygidial area. Fig. 63—66. Psenulus oweni sp. nov, 2, holotype. 63, thorax,

dorsal aspect; 64, propodeum, anterior aspect; 65, pygidial area; 66, face. Fig. 67—68. Psenulus
oweni sp. nov., &, paratype. 67, genitalia, dorsal aspect; 68, apex lateral valves

J. P. vAN LITH: Revision of Ethiopian Psenini 87

P. nigeriae is closely related to P. bidentatus (Cameron) and most resembles the form
provisionally considered to be the male of P. bidentatus pallidus, which is paler than the
female of P. bidentatus pallidus but still somewhat darker than P. nigeriae.

Psenulus ghesquierei Leclercq
Leclercq, 1961a: 34, 9 (Zaire).

New record: Uganda: 1 9, Bwamba, Hakitengya, Febr — March, 1949, Fan trap, coll.
W. H.R. Lumsden (BM).

I have compared this specimen with the holotype from Eala (Zaire), June, 1935
(MRAC) and consider them to be completely identical. As Leclercq has only given a
short characteristic of P. ghesquierei in his key (1961a: 19), a detailed redescription of
the holotype follows.

Female. — Head and thorax black; mandibles yellow with reddish-brown tips, palpi
testaceous, scapes yellow, second segment yellow with brown mark above, following
segments reddish-brown below, segments 3—9 brownish at apices, dorsally dark brown.
Median part of pronotum yellow dorsally, lateral corners black. Pronotal tubercles yellow.
Metanotum yellow, laterally reddish-brown. Tegulae yellowish, transparent. Veins of
wings dark brown, paler at base. Fore and mid legs including trochanters yellow, femora
darkened below, especially mid femora. Hind trochanters yellowish, hind femora brown,
hind tibiae reddish above, brown below, tarsi brownish. Petiole black, ventral plate of
petiole and gaster yellowish-red.

Frontal carina sharp, raised part between antennae not broadened, ending below in a
distinct transverse, bilobed, carina. Frons below anterior ocellus obliquely striato-punctate,
rest of frons and vertex sparsely finely punctate. Face and clypeus dull, protruding
median part of clypeus with two distinct teeth, distance between tips of teeth about one-
seventh of total distance there between the eyes. Tempora with fine longitudinal striation.
Mandibles normal. Third antennal segment about two and a half times, segments 4—5
about twice, sixth segment about one and a half times, segments 7—8 about one and
one-third times as long as broad at apex, segments 9—11 about as long as broad at
apex, last segment about twice as long as broad at base.

Scutum transversely rugoso-punctate, not very coarsely, punctures fine with shining
interspaces. Prescutal sutures indistinct, about as long as anterior third, parapsidal sutures
distinct. Scutellum and metanotum with sparse punctation. Propodeal enclosure triangular,
lateral parts with about six oblique carinae, central part with indistinct median line, upper
part of propodeum laterally with oblique striae, on both sides of median sulcus a small
smooth area. Sulcus broad. Back-side of propodeum coarsely reticulate, sides less coarsely
sculptured, dull anteriorly. Metapleura smooth on fore part, posterior triangular part with
a few oblique carinae. Mesopleura and mesosternum with superficial irregular punctation.
Anterior oblique suture gradually widening upwards, with long transverse carinae.
Anterior plate of mesepisternum with some irregular reticulation. Legs normal; mid
tibiae somewhat thicker at apex, outer side flattened and bordered behind by a bent row
of five short yellowish-red thorns. First recurrent vein of fore wings ending in second
submarginal cell, second recurrent vein ending in third submarginal cell. Upper side of
second submarginal cell about half as long as lower side. Petiole cylindrical, about one
and a half times as long as first tergite, rounded above, small depression at apex. Gaster

88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

very finely punctate, no distinct pygidial area. Base of second sternite without well-
defined depression.

Face with appressed silvery pubescence, head and thorax whitish pubescent, rather
dense and long on propodeum, yellowish on legs and gaster, apical margins of fourth
and fifth sternites with fringe of very short pale hairs, sixth sternite with dense yellowish-
golden pubescence.

Length at least 7.5 mm.

Male unknown.

Psenulus trevirus Leclercq
Leclercq, 1961a: 34, 4 (Zaire).

Redescription. Male. — Head and thorax black, with following yellow marking:
mandibles except reddish tips, palpi, pronotum dorsally and pronotal tubercles, two
square marks on scutellum, extreme upper corner of anterior plate of mesepisternum,
metanotum, with a narrow interruption, two marks on back of propodeum close to
longitudinal sulcus, as long as apical half of propodeum and each about one-fourth of
width of back of propodeum. Yellow colour on thorax somewhat whitish. Underside of
antennae yellowish-red. Fore and mid legs including trochanters yellowish, base and back
of fore femora and back of mid femora brown, hind trochanters and femora dark brown,
rest of hind legs reddish with median part of tibiae somewhat brownish. Petiole including
ventral plate black but extreme base of petiole below yellowish-white; first and base of
second tergite black, rest of gaster dark reddish, apex more brown. Sternites partly
darkened. Veins of wings brown.

Raised part of frontal carina not broadened between antennae, transverse carina below
antennae much protruding. Frons and vertex finely sparsely punctate. Clypeal margin
distinctly bidentate. Tempora smooth. Antennae long, filiform, third segment about two
and one-third times, segments 4—12 about twice as long as broad at apex, last segment
about two and a half times as long as broad at base, segments slightly convex below; no
tyloidea.

Anterior lateral corners of pronotum obtuse. Scutum coarsely punctate, interstices
irregular, prescutal sutures about one-third of length of scutum, indistinct, indicated by
transverse short grooves. Scutellum and metanotum almost impunctate. Enclosed area of
propodeum concave, triangular, with a few indistinct oblique carinae. Back of propodeum
with coarse reticulation, a smooth area behind lateral parts of enclosed area. Median
longitudinal sulcus neither broad nor narrow. Sides of propodeum except narrow anterior
part with coarse reticulation. Mesopleura distinctly punctate, interstices about three or
four times size of punctures, hypo-epimeral area almost impunctate. Anterior oblique
suture foveolate, widened upper part smooth. Legs normal. Base of second submarginal
cell about one and a half times as long as upper side. Petiole long and cylindrical, about
one and one-third times as long as first tergite in dorsal view, first tergite about one and
a half times as long as wide at apex, slightly nodose. Petiole dorsally rounded, with small
apical pit. Apex of gaster laterally depressed.

Face with appressed yellowish-silvery pubescence. Pubescence of thorax greyish, of
gaster yellowish, petiole with long erect lateral hairs.

Length about 8 mm.

J. P. VAN LITH: Revision of Ethiopian Psenint 89

Female unknown.

Material studied: Zaire, “Equateur: Bokuma, II—1952, R. P. Lootens”, 1 ¢ (holo-
type) (MRAC).

The systematic place of this species is still unclear. The shape of the antennae does not
point to a close relationship to P. bidentatus (Cameron), as Leclercq suggested (19612).
It may be more closely allied to P. oweni sp. nov., from Sierra Leone.

Psenulus oweni spec. nov.

Female. — Head black; mandibles pale yellow with reddish-brown tips, labrum brown,
palpi pale yellow. Scape of antennae pale yellow, extreme tip brown, following antennal
segments dark brown, almost black dorsally, underside reddish-yellow, tip of last segment
brownish below. Thorax black with following yellow parts (Fig. 63): dorsum and upper
margin of foreside of pronotum, pronotal tubercles, longitudinal mark along tegulae, two
narrow marks on scutum, widened towards apex, commencing at end of prescutal sutures
and ending well before apical margin of scutum — marks on scutum, especially the two
median ones, may be more or less reduced, rarely lacking — a small mark on axillae —
often missing — scutellum including lateral margin behind depressed lateral part,
posterior margin sometimes darkened medially, metanotum with sometimes medially
darkened posterior margin, back of propodeum, except two black triangular incisions
projecting from the black base (Fig. 64) and black apical margin, upper third of anterior
plate of mesepisternum, and a small mark on hypo-epimeral area, which may be lacking.
Fore and mid legs including trochanters and tip of coxae yellowish-white, back of mid
tibiae and mid tarsi somewhat orange; hind coxae pale yellow with black base, rest of
hind legs reddish-orange, back of trochanters brownish, femora sometimes with brown
streak behind, tarsi light-brown. Tegulae yellowish-red, veins of wings blackish-brown.
Petiole black, basal third yellowish, gaster including ventral plate of petiole reddish.

Protruding median part of clypeal margin weakly emarginate (Fig. 66). Frontal carina
distinct, raised part between antennae narrow, slightly flattened, ending below antennae
in a small shining point; no distinct transverse carina, at most slightly indicated. Frons,
vertex and tempora shining, almost impunctate. Occipital carina ending in hypostomal
carina. Mandibles normal. Third antennal segment just over two and a half times, fourth
segment over twice, fifth segment about twice, segments 6—9 about one and a half times,
tenth segment about one and a third times and eleventh segment about one and a quarter
times as long as broad at apex, last segment about one and a half times as long as broad
at base. Pronotal corners with sharp angle; scutum densely, extremely finely punctate
with a few larger punctures. Prescutal sutures about half as long as scutum. Scutellum
and metanotum very finely punctate. Propodeal enclosure triangular, hind margin not
sharply defined, median two longitudinal carinae slightly bent inwards at base, lateral
parts with oblique carinae (Fig. 64). Median longitudinal suture on back of propodeum
very narrow, ending in a shallow depressed triangular area at apex of propodeum. Back
of propodeum smooth, dorso-laterally with weak parallel striae, sides of propodeum and
metapleura smooth. Mesopleura and anterior plate of mesepisternum and mesosternum
with fine hair-bearing punctures. Anterior oblique suture narrow, with a few transverse
carinae, widened upper part smooth. Epicnemial areas shining. Legs normal, mid tibiae
postero-dorsally with three or four reddish spines, base of hind tibiae on outer side with
short flattened area, margined basally by a semi-circular area of small and short reddish

90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

spines. Upper side of second submarginal cell nearly two-thirds of length of lower side,
first recurrent vein ending distinctly in first submarginal cell, second recurrent vein
ending well in third submarginal cell. Petiole in dorsal view about one and one-third
times as long as first tergite, cylindrical, apex dorsally with small triangular pit. Gaster
slender, almost impunctate, first tergite somewhat nodose, pygidial area distinct, narrow
elongate-triangular (Fig. 65). Base of second sternite somewhat depressed.

Lower part of frons and face with appressed silvery pubescence, clypeus also with long
erect hairs. Vertex and tempora with short silvery pubescence, vertex also with long hairs.
Thorax with whitish pubescence, on scutum short, intermixed with long hairs. Pubes-
cence long and denser on scutellum, metanotum and propodeum, very dense on mesoster-
num. Epicnemial areas below with patch of dense appressed yellowish pubescence.
Pubescence of legs and gaster yellowish, hind margins of fourth and fifth sternites with
dense and short yellowish-white pubescence, sixth sternite with large patch of yellowish-
golden pubescence, petiole with long outstanding hairs in two lateral rows on dorsum and
on sides.

Length about 8—9 mm.

Male. — Similar to female, with following differences. Small yellow mark on hypo-
epimeral area lacking, yellow markings on scutum completely lacking in the allotype,
lateral hind margin of scutellum behind depressed lateral area sometimes black. Yellow
marking on back of propodeum reduced to four marks, more or less united below. Fore
and mid coxae brown, hind coxae black at base, posterior half reddish; fore and mid
tibiae with brown streak on back, hind femora dark brown, underside reddish, hind
tibiae dark brown on inner side, base reddish-brown. First gastral tergite somewhat
brown at base and sternites 3—4 and 6 more or less dark brown.

Face with distinct transverse carina below antennae. Antennae long, segments distinctly
convex below, segments 3—12 slightly more than twice as long as broad at apex, last
segment nearly three times as long as broad at base. Genitalia: Fig. 67,68.

Length about 8—9 mm.

Sierra Leone: Freetown, 1 9 (holotype), Febr., 1968, 1 g' (allotype), Jan., 1968;
19, Jan, 1967, 1 2, March, 1967,01 2, April, 1967, 6 9, May, 1967, /2\9*May,
1968, 1 &, March, 1967, 1 &, Dec., 1967 (all paratypes), coll. D. F. Owen (HT, a
few paratypes in author’s collection).

Only in the holotype and in the allotype the marks are fully yellow, in the paratypes
the colour is more whitish, the chitin being also partly transparent there.

P. oweni is easily distinguished from P. nigeriae by the lacking transverse carina below
the antennae of the females and the richer yellow marking. It shows some relationships
with the group of P. capensis. The male much resembles P. trevirus from Zaire.

Psenulus dilectus (Saussure)

Saussure, 1892: 571—572, Q (Psen dilectus; Malagasy: province of Imerina).

Schulz, 1911: 141—142 (Psenulus dilectus (Saussure) = Stenomellinus dilectus Saussure in litt.).
Arnold, 1945: 154, 2 (Malagasy: Bekily).

Leclercq, 1961a: 35.

Leclercq, 1961c: 117, @ (Malagasy: Sakavondro, Ranomafana).

Material studied: 1 9, “Madagascar”, det. J. Leclercq (FAG); 1 9, “Madagascar,
Bekily, Reg. sud de l’île”, Oct., 1936, coll. A. Seyrig (RM).

J. P. vAN LITH: Revision of Ethiopian Psenini 91

This species resembles those placed in the group of P. capensis Brauns in many respects;
especially the vertical carinae below the antennal sclerites and the nodose first gastral
tergite are striking features. However, P. dilectus is less slender, the transverse carina
below the antennae is distinct, though very fine, and it has a distinct, narrow, pygidial
area; the colour is also different. The male has not yet been found.

A few characters should be added to earlier descriptions. — Median part of clypeal
anterior margin protruding, weakly emarginate. Occipital carina ending normally in
hypostomal carina. Pronotal corners obtuse. Prescutal sutures distinct only on basal third
of scutum. Propodeum behind enclosed area smooth and shining, backside sparsely finely
punctate, apex with some reticulate carination. Petiole cylindrical, apex dorsally with
small triangular pit, petiole slightly longer than first tergite, which is about twice as
long as wide at apex and distinctly nodose. Apical margins of fourth and fifth gastral
sternites with dense short pubescence.

Psenulus uelleburgi spec. nov.

Female. — Head black; mandibles yellow with dark. tips, labrum yellowish-red, palpi
straw-yellow, scape of antennae yellowish with reddish tip, following segments yellowish-
red below, dorsally dark brown, last segment dorsally reddish-brown. Thorax black, with
following parts (Fig. 69) yellow: dorsal side and upper half of foreside of pronotum,
pronotal tubercles, large yellow mark on sides of scutum reaching fore margin, two
narrow lines along prescutal sutures, about half as long as scutum, posterior half of
axillae, scutellum and metanotum including posterior lateral margins, propodeum except
upper lateral corners and apex, round yellow mark on anterior half of hypo-epimeral
area, anterior plate of mesepisternum. Lower half of foreside of pronotum and greater
part of mesopleura reddish. Fore legs including coxae yellow, foreside of mid legs yellow,
back side of mid legs mostly brownish-red, hind legs reddish-brown including tibial
spurs, apical half of hind femora yellowish below. Basal third of petiole yellow, rest dark
brown above, ventral plate yellowish. Gaster brown, darkening towards apex. Veins of
wings dark brown. Tegulae yellowish-red.

Raised part of frontal carina between antennae narrow, carina ending below antennae
in a fine transverse carina, which is about half as long as distance between eyes.
Supraclypeal area and upper half of clypeus dull, very finely punctate, anterior half
shining, protruding median part of clypeus slightly emarginate, frons and vertex shining.
Tempora smooth and shining. Occipital carina ending normally in hypostomal carina.
Apex of mandibles normal. Antennae long and slender, third segment about three times,
segments 4-5 about two and a half times, sixth segment about twice, segments 7—8
about one and three quarters, segments 9—11 about one and a half times as long as broad
at apex, last segment about two and a half times as long as broad at base.

Pronotal corners with obtuse angle. Scutum shining, sparsely distinctly punctate.
Prescutal sutures fine, about one-third of length of scutum. Parapsidal sutures sharp,
linear. Enclosed area of propodeum triangular with oblique carinae on lateral parts, these
parts not sharply separated from rest of propodeum, median part with irregular fine
median carina. Upper third of back of propodeum smooth, finely obliquely striate below,
lower two-thirds and sides of propodeum finely reticulate, fore part of sides smooth.
Median longitudinal groove on back of propodeum narrow. Metapleura smooth and
shining, mesopleura and mesosternum smooth with fine punctures. Anterior oblique

92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

suture narrow, indistinctly foveolate, widened upper part smooth. Foreside of fore femora
much flattened; apical two-fifths of mid tibiae with row of four dark red thorns; hind
legs slender, base with small smooth depressed area, margined at base by semicircular
row of thin reddish teeth. First recurrent vein of fore wings interstitial, second recurrent
vein ending just in third submarginal cell. Upper side of second submarginal cell about
two-thirds of length of lower side. Petiole about twice as long as first tergite, cylindrical,
apex with relatively large triangular pit. Gaster elongate, first tergite much convex,
pygidial area narrow and shining, with long parallel lateral carinae (Fig. 70).

Pubescence of head silvery, on face below antennae mostly appressed, yellowish on
thorax, dense and silvery on mesosternum, epicnemial areas below with round patch of
dense yellowish pubescence, last sternite densely yellowish-golden pubescent. Petiole with
few long, obliquely downwards directed hairs.

Length about 10.5 mm.

Male unknown.

Equatorial Guinea (Spanish Guinea): 1 9 (holotype), “Westafrika, Uelleburg”,
June—Aug., 1908, coll. Tessmann (ZMB).

P. nelleburgi seems to be rather closely related to P. dilectus from Malagasy.

Psenulus capensis capensis Brauns

Brauns, 1899: 391—392, Q and & (Cape Province near Port Elizabeth).

Arnold, 1923: 47—48, & (Rhodesia: Matopos).

Arnold, 1947: 159—160, 9 and 4 (Psenulus pauxillus sp. nov.; Zambia: Abercorn).
Leclercq, 1961a: 30—32 (Zaire; S. Africa: Natal, S. Zululand; Cape Province).
Leclercq, 1961b: 56—57 (Zaire). i

Arnold, 1962: 851-852, 2 (Psenulus stuckenbergi sp. nov., Mozambique: Marromeu).

New records: South Africa: Natal: “Mfongosi, Zululand” (Mfongozi, near Krans-
kop), 1 g', Febr., 1914, 1 9, May, 1916, coll. W. E. Jones (SAM); 1 g', Durban, no.
4457 (AMNH).

South Africa, Transvaal: Ellisras, N. Transvaal, 2 9, 6 April and 19 June, 1962, 3 9,
22 June, 1963-1 O13 May 972 378. 19) June, 1962, 278, (220]june MIO OSE ME
Schoemanville, H-B Dam, 8 Aug., 1965, all coll. H. N. Empey (EC, one 9 and one
d' PMFV); 1 9, 5 miles N of Warmbad, 24 Febr., 1968, coll. Krombein and Spangler
(USNM).

Zaire (Congo): 2 9, Kasongo, Aug., 1959, coll. P. L. G. Benoit, 2 G', Kivu: Uvira,
Sept., 1958, coll. J. Pasteels (MRAC); 1 9, Tshibinda, Aug., 1931, coll. J. Ogilvie,
1 G', Lubumbashi (Elisabethville), Sept., 1931, coll. L. Ogilvie (BM).

Mozambique: 1 g', Beira, Dec., 1960 (RM).

Tanzania: 1 G', “Tanganyika, Kilosa”, 25 Aug., 1948, at flowers of Solanum incanum,
coll. G. Salt, BM 1971—85 (BM).

Malawi: “Nyassa-See Langenburg”, 1 g', 31 Oct—14 Nov., 1898, 1 g', end Dec.
1898—end Jan., 1899, 3 9, 17 Aug., 1899, all coll. Fülleborn (ZMB).

Uganda: 1 ©, “Buunga’” (?), 29 May, 1924, coll. G. L. R. Hancock, pres. by Com.
Inst. Ent. BM 1973—1 (BM); 1 ©, Ankole, Kichwamba, 23—29 April, 1968, coll.
P. J. Spangler (USNM).

Sierra Leone: Freetown, 1 ©, July, 1967, 4 &, March and July, 1967, Jan. and Febr.,

J. P. vAN LITH: Revision of Ethiopian Psenini 93

1968, coll. D. F. Owen (HT); 1 9, “Rokupr.” (?), Febr., 1965, CIE 612 (241), pres.
by Com. Inst. Ent. BM 1967—3 (BM).

In one of the two females from Kasongo (Zaire) the fore tibiae are completely
yellowish-white, the females from Malawi have a brownish streak on the back of the
fore tibiae.

P. capensis Brauns and P. lattannulatus (Cameron) are distinctly different species, the
former having a pronotum with obtuse lateral corners (Fig. 74), whilst in P. Jatiannu-
latus the lateral corners always have a more or less sharp angle (Fig. 71 and 72). More-
over P. capensis is smaller, length in both sexes being about 7 mm (in P. lattannulatus
about 8—9 mm), and the sculpture on frons, scutum and propodeum is generally much
coarser.

The female of P. capensis is easily recognized by the traces of a transverse carina below
the antennae, consisting of a short vertical carina downwards from each of the antennal
sclerites, best visible in dorsal view, when it looks like a tooth. They are connected by a
more or less distinct transverse low ridge, never forming a sharp carina. There is no
distinct pygidial area but sometimes two very close parallel carinae are indistinctly
indicated. Upper part of back of propodeum usually distinctly obliquely striate.

The antennae of the male have low but distinct narrow-elliptic tyloidea, small on
fourth segment, on segments 5—10 about half as long as segments, smaller on segments
11—12, sometimes very indistinct on twelfth segment. Dorsal side of flagellum blackish-
brown, lower side yellowish-brown; tyloidea situated on back-side of segments just in the
dark part but they are somewhat paler brown. P. lattannulatus has no tyloidea.

I did not see Braun’s types but his excellent description allows to recognize his species
without any doubt. He labelled the female and the male from “Algoa Bay, Capland”,
as “Q type” and “g' type”; I have designated the female as the lectotype.

I have seen the type of P. stuckenbergi Arnold (1962), a female without gaster, and
consider this a true P. capensis capensis. Arnold’s figure No. 6 of the face of P. stucken-
bergi is not exact.

P. capensis is one of the very few Ethiopian Psenini, of which we know some partic-
ulars in regard to its biology. Brauns found his couple in the month of August in stems
of a species of Phragmites in which he thought this Psenulus should be nesting. Mr.
Empey (letter of 23 Nov., 1972) collected his specimens about the middle of June — in
the South African winter — in the subtropical area of North Transvaal (Ellisras),
hovering about the reeds on the banks of the river. Finally, the female from “Rokupr.”,
Sierra Leone, caught in February, bears on its pin the remains of its cocoon. This cocoon
very much resembles that of the Palaearctic P. schencki Tournier. It has a solid, flat,
upper side, smooth and creamy-white on its lower surface, with remains of the inner wall
of the stem solidly fastened to its circumference. The sides and bottom of the cocoon are
made of a fine whitish tissue, probably covering the whole length of the cell, but keeping
free from its wall.

This specimen from Sierra Leone also bears the following note on its pin: “pred. on
Hecalus on wild rice”. Hecalus is a Homopteron belonging to the Cicadelloidea (or
Jassoidea), Hecalidae. Jassids as well as Delphacids are known as the prey of Indo-
Australian Psenulus. The food of Psenulus sogatophagus Pagden consists of the Delphacid
Sogata furcifera Horv. and the Jassid Nephotettix bipunctata F., both Homoptera being
pests of rice (Pagden, 1933).

94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

Psenulus capensis laevior Arnold

Arnold, 1951: 163, 4 (Psenulus pauxillus laevior; Ethiopia).
Leclercq, 1961a: 30—31 (Psenulus capensis).

Arnold distinguishes this form from P. pauxillus Arnold (= P. capensis Brauns) by
the brown colour of the femora which are black in P. capensts, the somewhat wider
scutum, the lacking feeble transverse ridges between punctures of scutum and the finer
sculpture of dorsum of propodeum (epinotum). Because of the variation in the sculpture
of P. capensis series of both sexes of the Ethiopian form should be studied to place it
with more certainty.

Psenulus latiannulatus latiannulatus (Cameron)

Cameron, 1910: 281—282, “9” (= 4) (Psen latiannulatus; Tanzania: Kilimanjaro)
Turner, 1912: 413, & (Nigeria).

Arnold, 1923: 50.

Leclercq, 1961a: 30—32, 9 and Â (Zaire).

Leclercq, 1961b: 57 (Zaire).

New records: Uganda: 1 g', Ankole, Kichwamba, 23-29 April, 1968, coll. P. J.
Spangler (USNM).

Rhodesia: 1 9, Cashel, 6 Dec., 1945 (RM).

Zaire (Congo); 1 g', Boma, 13°0’ E, 6°0’ S, 14 June, 1915, coll. Lang and Chapin
(AMNH); 1 9, Kasongo, Sept., 1959, coll. P. L. G. Benoit (MRAC), 1 g', Lubum-
bashi (Elisabethville), 25 Sept., 1962, coll. M. and A. and J. Bourgeois (HT).

Nigeria: 1 g', Aba, May, 1967, coll. Jean Townes (HT).

Sierra Leone: Freetown, 8 9, March, May, July, Oct., Dec., 1967, Febr., April, 1968,
12 g', Jan, July, Aug., Sept., Oct., Dec., 1967, Jan., 1968, all coll. D. F. Owen (HT).

Dr. Per Inge Persson of the Naturhistoriska Riksmuseet at Stockholm kindly sent me
the holotype, which is labelled: “Kilimandjaro, Sjöstedt. 1905—6, Kibonoto, 1300—
1900 m, 4 maj, Typus, Psen latiannulatus Cam. Type (in Cameron’s writing). 307.3”.
From Cameron’s description of the face of his type it is evident that he had a male before
him. This has been proved by examination of the holotype, so Cameron has again been
misled by the long apical spine of the males of the Psenini.

The anterior corners of the pronotum are almost rectangular in the type (Fig. 72), not
as sharp as in other P. lattannulatus studied (Fig. 71, male from Sierra Leone), but
certainly not as obtuse as in P. capensis. Upper side of antennae dark brown, underside
paler brown, more reddish at base. Although the antennae are somewhat dirty, it is
sufficiently clear that there are no tyloidea. Upper part of propodeum behind enclosed
area close to median longitudinal suture almost smooth, behind this smooth part a few
superficial oblique rugae, back coarsely reticulate.

Fig. 75 shows sternites 5—8 of a male from Sierra Leone, somewhat extracted and
bent upwards, in lateral view. Apical margin of seventh sternite (Fig. 76) not emarginate.
Genital apparatus slender (Fig. 77, 78).

Pygidial area of female indistinct, in some specimens last tergite with two very super-
ficial, very close and parallel ridges. Apical margins of fourth and fifth sternites with
dense short pale pubescence.

There is some variability in the angle of the pronotal corners as well as in the puncta-

J. P. vAN LITH: Revision of Ethiopian Psenini 95

Fig. 69—70. Psenulus uelleburgi sp. nov. ©, holotype. 69, thorax; 70, pygidial area. Fig. 71.

Psenulus latiannulatus latiannulatus (Cameron), &, Sierra Leone. Pronotum. Fig. 72. Psenulus

latiannulatus latiannulatus (Cameron), 4, holotype. Right part pronotum. Fig. 73. Psenulus latian-

nulatus basilewskyi Leclercq, 6. Pronotum. Fig. 74. Psenulus capensis capensis Brauns, 6, North

Transvaal. Pronotum. Fig. 75—78. Psenulus latiannulatus latiannulatus (Cameron), 6, Sierra Leone.

75, 5th, 6th, 7th and 8th sternites, lateral aspect, 7th and 8th sternites bent upwards; 76, 7th sternite,
lateral aspect; 77—78, genitalia, dorsal and ventral aspect

96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

tion of the thorax. The legs of the specimens from Sierra Leone are usually more whitish
than in specimens from other regions. The female from Cashel, Rhodesia, bears the
following note, probably in Arnold’s handwriting: “very near Psenulus lattannulatus, but
2nd abs. radius differs”.

Length of female about 8—9 mm, of male about 8 mm.

P. latiannulatus belongs to a small group which is easily recognized by the slender
form and the nodose first gastral tergite. P. latiannnlatus and its nearest relative, P.
capensis, have a wide area of dispersal over the Ethiopian region. P. dilectus Saussure
may be a representative of the group in Malagasy, but the female of this species has a
distinct narrow pygidial area.

Psenulus latiannulatus basilewskyi Leclercq

Leclercq, 1955: 420—421, 4 (Psenulus basilewskyi, Rwanda).
Leclercq, 1961a: 32—33, 9 and 4 (Psenulus capensis basilewskyi; Zaire).

New records: Cameroon: 1 ®, Yaoundé “Amt Jaunde, Buschwald”, 19—23 Oct.,
1914, coll. Tessmann (ZMB). Length, sting not included, about 7.5 mm.

Zaire (Congo): 1 ©, Kasongo, Sept., 1959, coll. P. L. G. Benoit (MRAC).

Further material studied from Zaire: 1 9 (allotype), Kiamokoto-Kiwakishi, Parc de
l’Upemba, 1070 m, 4—16 Oct., 1948 (MRAC); 1 & (paratype), Eala (Equateur),
Nov 1931, det..J. Leclercq) (FAG):

In the allotype ( © ) only the basal half of the antenna is yellowish-red, rest of under-
side dark brown, dorsal side black. Apex of fore femora on foreside, base and foreside
of fore tibiae yellow, basitarsi yellowish, following segments of tarsi brownish. Base of
mid tibiae yellow on foreside, tarsi including basal segment dark brown. The female from
Cameroon has also foreside of mid tibiae somewhat brownish. In the female from Kason-
go apical half of fore femora, fore tibiae completely, and basal two-thirds of foreside of
mid tibiae yellowish-white. Antennal segments 4—12 in this specimen with reddish-
brown mark below.

Propodeum behind enclosed area and back of propodeum of female smooth, with very
fine dorso-lateral striae.

Male without distinct tyloidea.

Length of female about 7.75 mm, of male about 7.5-mm.

The sharp anterior corners of the pronotum (Fig. 73) are an indication of the close
relationships between P. basilewskyi and the nominate form of P. lattannulatus (Came-
ron). P. basilewskyi is easily distinguished in both sexes by the dark base of the hind
tibiae. The apex of the mid tibia is more or less brownish. The superficial close, parallel
carinae on the pygidium of the female are slightly more distinct than in P. lattannulatus
s.str.

REMARKS ON RELATIONSHIPS

Psen Latreille

Psen silvaticus Arnold (Rhodesia) and Psen leclercgi sp. nov. (Malagasy) probably
belong to the subgenus Psen. Both species are remarkable because of the coarse and
dense sculpture of their mesopleura. The males, still unknown, would be welcome to

J. P. VAN LITH: Revision of Ethiopian Psenini 97

confirm the subgeneric status. Psen congolus Leclercq (Zaire) certainly belongs to Psen
s.str. as is also proved by the sternal tufts of hair in the male.

Psen madecassus Arnold (Malagasy) differs from Psen s.str. in the sculpture of the
petiole and the absence of fasciculate hairs on the sternites of the male. It may belong to
the subgenus Punctipsen, known from the palaearctic and oriental regions. __

Psen patellatus Arnold (South-West Africa) is another species difficult to place.
Although the antennae resemble those of Psen ater F., it differs in many respects so much
from other Psen s.l. that the erection of a new subgenus seems to be justified. We hope,
however, that soon the female will be discovered (the males date from half a century
ago!).

Psen empeyi sp. nov. (South Africa) is a distinct representative of the subgenus
Mimesa.

Psenulus Kohl

Most of the Ethiopian species of Psenulus closely resemble their Indo-Australian
relatives. In the majority of species the interantennal carina is broadened (primitive
character?) as in all but one of the palaearctic species and about one third of the oriental
forms. On the other hand the petiole is cylindrical or nearly so, which means that it is
more evolved. The dorsal groove of the petiole is lacking, the sides of the first sternite,
out of which the petiole has developed, being completely merged. It is already possible to
distinguish one large group and a few smaller groups of closely allied species.

The group of Psenulus turneri Arnold is characterized by the coarse sculpture of the
tempora and in the females by the fringes of long hairs on the fourth and fifth gastral
sternites. It has this combination of characters in common with the group of Psenulus
rugosus from the Philippines and Borneo. Also the triangular second submarginal cell of
the oriental species is found in some of the African forms, where in some cases the cell
is even petiolate. The Ethiopian group comprises turneri Arnold (South Africa), paulisae
Leclercq, erusus Leclercq and ealae Leclercq (Zaire and Sierra Leone), garambae Leclercq
(Zaire), aurifasciatus sp. nov. (Sierra Leone), freetownensis sp. nov. (Sierra Leone and
Equatorial Guinea), patei Arnold and saltitans Arnold (Rhodesia), probably also 4/veo-
latus sp. nov. (Rhodesia) and a related, still unnamed male from Angola.

Psenulus reticulosus Arnold (Malagasy) and sapobaensis sp. nov. (Nigeria), with
rather long petiole, oblique striae on the back of the propodeum and indistinct or absent
pygidial area in the female may be closely related.

Group of Psenulus luctuosus Arnold. This seems to be a group distinguished by the
broadened part of the interantennal carina which usually ends with two parallel carinae
in the transverse carina. The pygidial area of the female is distinct. The following species
should be included: /ct4osus Arnold and stevensoni Arnold (Rhodesia), jacoti sp. nov.
(South Africa), lwsingae Leclercq (Zaire) and rugifrons sp. nov. (Ethiopia), probably
also avernus Leclercq (Zaire).

In the group of Psenulus bidentatus (Cameron) not only the nominate form from
Tanzania, Kenya, Rhodesia, Zaire and Angola, the subspecies rabrocaudatus Turner
(South Africa) and pallidus subsp. nov. (Zaire, Rhodesia, Malawi and Equatorial
Guinea) can be placed, but also nigeriae Leclercq (Nigeria, Sierra Leone).

A peculiar group is formed by the widely distributed Psenulus capensis Brauns and its
near relatives lattannulatus (Cameron) with a somewhat more restricted area, latiannula-

98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

tus basilewskyi Leclercq (Zaire, Rwanda, Cameroon), dilectus (Saussure) (Malagasy) and
perhaps also zelleburgi sp. nov. (Equatorial Guinea) and oweni sp. nov. (Sierra Leone).
They all have a strongly or slightly nodose first gastral tergite, the raised part of the
carina between the antennae is narrow, the transverse carina below the antennae is fine
or indistinct in the female; the pygidial area of the female is indistinct in capensis and
latiannulatus, distinct elongate-triangular in the other forms which probably belong to
this group. The Indo-Australian group of Psenulus pulcherrimns, especially its subgroup
of carinatus, has the following characters in common: more or less nodose first gastral
tergite, narrow interantennal carina and indistinct transverse carina below antennae in the
female, but in that group the thorax is yellow marked and the pygidial area of the female
is distinct.

Psenulus benoiti Leclercq (Zaire) is very remarkable, the pygidial area of the female
being reduced to a single sharp and long median longitudinal carina. So far this character
was only known of a few Mexican species.

The relationships of the remaining species are not clear at the moment. For this reason
the species have been listed hereafter in alphabetical order.

PROVISIONAL LIST OF PSENINI OF THE ETHIOPIAN REGION INCLUDING MALAGASY

Genus Psen Latreille

Subgenus Psen Latreille

P. silvaticus Arnold, 1924, 9 — Rhodesia
P. leclercgi sp. nov., 2 — Malagasy
P. congolus Leclercq, 2 and & — Zaire
Subgenus ?
P. madecassus Arnold, 1945, © and & — Malagasy
Subgenus ?
P. patellatus Arnold, 1924, & — South-West Africa
Subgenus Mimesa Shuckard
P. empeyi sp. nov, d — South Africa (Transvaal)
Genus Psenulus Kohl
P. alveolatus sp. nov., g' i — Rhodesia
P. aurifasciatus sp. nov., 9 — Sierra Leone
P. avernus Leclercq, 1961, 4 — Zaire
P. benoiti Leclercq, 1961, © and & — Zaire
P. bidentatus bidentatus (Cameron), 1910, 9 and & — Tanzania, Kenya, Rhodesia,
Angola, Zaire
P. bidentatus pallidus subsp. nov., © and & — Zaire, Rhodesia, Malawi,
Equatorial Guinea
P. bidentatus rubrocaudatus Turner, 1912, 9 — South Africa (Natal)

P. bidentatus subsp. 9 — Ethiopia

liae Nu Ik:

Vv PPP

BP,

ini u Bee Bye

J. P. van LITH: Revision of Ethiopian Psenini 99

capensis capensis Brauns, 1899, 9 and J

capensis laevior Arnold, 1951, &
dilectus (Saussure), 1892, 9
ealae Leclercq, 1961, 9 and 3
erusus Leclercq, 1961, 9
freetownensis sp. nov., 9 and g

fulgidus Arnold, 1945, 9 and &
garambae Leclercq, 1961, 9
ghesquierei Leclercq, 1961, 9
jacoti sp. nov., Q and G

kobli Arnold, 1923, 9
latiannulatus latiannulatus (Cameron), 1910,

Q and g°

latiannulatus basilewskyi Leclercq, 1955, 9 and G'
leoninus sp. nov., 9

luctuosus Arnold, 1929, 9
lusingae Leclercq, 1961, 9

nigeriae Leclercq, 1961, 9

oweni sp. nov., Q and g

patei Arnold, 1940, 9

paulisae Leclercq, 1961, 9 and g'
reticulosus Arnold, 1945, © and J
rugifrons sp. nov., 9

saltitans Arnold, 1958, 9
sapobaensis sp. nov., 2

stevensoni Arnold, 1940, 9 and J
trevirus Leclercq, 1961, G'

turneri Arnold, 1927, Q and J'

uelleburgi sp. nov. 9

— South Africa (Cape

Province, Natal, Transvaal),
Mozambique, Rhodesia,
Malawi, Zambia, Tanzania,
Uganda, Zaire, Sierra Leone
Ethiopia

Malagasy

Zaire, Sierra Leone

Sierra Leone, Equatorial
Guinea

Malagasy

Zaire

Zaire, Uganda

South Africa (Cape
Province, Transkei)
Rhodesia

Tanzania, Rhodesia,
Uganda, Zaire, Nigeria,
Sierra Leone

Zaire, Rwanda, Cameroon
Sierra Leone

Rhodesia

Zaire

Nigeria, Sierra Leone
Sierra Leone

Rhodesia

Zaire, Sierra Leone
Malagasy

Ethiopia

Rhodesia

Nigeria

Rhodesia

Zaire

South Africa (Cape
Province)

Equatorial Guinea

100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974

REFERENCES

Arnold, G., 1923. The Sphegidae of South Africa. Part IV. — Ann. Transv. Mus. 10 (1): 1—58.

1924. The Sphegidae of South Africa. Part V. — Ann. Transv. Mus. 11(1): 1—73.

1927. The Sphegidae of South Africa. Part VIII. — Ann. Transv. Mus. 12 (2): 55—131.

1929. The Sphegidae of South Africa. Part XIV. — Ann. Transv. Mus. 13 (4): 381—418.

1930. A checklist of the Sphegidae of the Ethiopian Region. — Cambridge, for The Trans-

vaal Museum, Pretoria, South Africa: 1—21.

1940. New Species of African Hymenoptera No. 4. — Ann. Transv. Mus. 20 (2): 101—

143.

1945. The Sphecidae of Madagascar. — Natn. Mus. S. Rhodesia: 1—193.

1947. New Species of African Hymenoptera No. 7. — Occ. Pap. natn. Mus. S. Rhodesia 13:

1312167.

1951. Sphecidae and Pompilidae (Hymenoptera) collected by Mr. K. M. Guichard in West

Africa and Ethiopia, 1941—1948. — Bull. Brit. Mus. (Nat. Hist.), (Ent.) 2 (3): 95—183.

1958. New Species of African Hymenoptera No. 13. — Occ. Pap. natn. Mus. S. Rhodesia

22B: 119—143.

1962. New Species of African Hymenoptera No. 16. — Occ. Pap. natn. Mus. S. Rhodesia

26B: 844—855.

Bohart, R. M. & E. E. Grissell, 1969. New Species of Psenini (Hymenoptera: Sphecidae). — Pan-
Pacif. Ent. 45 (3): 216—221.

Brauns, H., 1899. Zur Kenntnis der südafrikanischen Hymenopteren. — Annln naturh. Mus. Wien
13: 383—423.

Cameron, P., 1910. Sjéstedts Kilimandjaro-Meru Expedition, Stockholm. 8. Hymenoptera. 7. Fossores
2: 197—296.

Leclercq, J., 1955. Contributions à l'étude de la faune entomologique du Ruanda-Urundi (Mission
P. Basilewsky 1953) LXXVII. Hymenoptera Sphecidae. — Annls Mus. r. Congo Belge 40:
404—426.

—, 1960. Hyménoptères Sphécides de Madagascar. — Bull. Annls Soc. r. ent. Belg. 96: 96—
100.

—, 1961a. Psenini (Hymenoptera Sphecoidea) Pemphredoninae. — Explor. Parc natn. Upemba
Miss. G. F. de Witte 60 (3): 13—36.

——, 1961b. Sphecoidea: Sphecidae Subfam. Sphecinae, Pemphredoninae et Crabroninae. —
Explor. Parc natn. Garamba Miss. H. de Saeger 20 (3): 43—105.

———, 1961c. Hyménoptéres Ampulicides et Sphécides récoltés par le Dr. Fred Keiser à Madagas-
car. — Verh. naturf. Ges. Basel 72: 100—119.

Malloch, J. R., 1933. Review of the Wasps of the Subfamily Pseninae of North America (Hymeno-
ptera: Aculeata). — Proc. U.S. natn. Mus. 82: 1—60.

Pagden, H. T., 1933. Two new Malayan Sphecoids. — Trans. R. ent. Soc. Lond. 81: 93—101.

Saussure, H. de, 1892. Histoire naturelle des Hyménoptères, in A. Grandidier, Hist. phys. nat. et
pol. de Madagascar, Paris 20: XXI + 1—590.

Schulz, W. A., 1911. Zweihundert alte Hymenopteren. — Zool. Ann. 4: 141—142.

Turner, R. E., 1912. Notes on Fossorial Hymenoptera. — Ann. Mag. nat. Hist. (8) 9: 410—423.

J. P. VAN LITH: Revision of Ethiopian Psenint

INDEX

The names of new species and subspecies are marked with an asterisk

alboscutellatus 70
alveolatus 68, 97
ater 97
aurifasciatus 65, 97
avernus 77, 97

basilewskyi 96, 98
benoiti 73, 98
bequaerti 82
bidentatus 81, 97

capensis 92, 97
carinatus 98
congolus 53, 97

dilectus 90, 98
ealae 64, 97
empeyi 58, 97

erusus 62, 97

freetownensis 67, 97
fulgidus 69

garambae 62, 97
ghesquierei 87

se
Ed

jacoti 77, 97
kohli 74

laevior 94
latiannulatus 94, 97
leclercqi 53, 96
leoninus 72
luctuosus 75, 97
lusingae 76, 97
luteopictus 73

madecassus 56, 97
mayorum 73
Mimesa 58, 97

nigeriae 85, 97
oweni 89, 98

pallidus 83, 97
patei 63, 97
patellatus 58, 97
paulisae 61, 97
pauxillus 92, 94

Psen 41, 96
Pseneo 58
Psenulus 42, 97
pulcherrimus 98
Punctipsen 56, 97
pygidialis 59

reticulosus 70, 97

rubrocaudatus 81, 84, 97
* rugifrons 80, 97

rugosus 97

saltitans 63, 97
sapobaensis 70, 97
schencki 93
silvaticus 52, 96
sogatophagus 93
Stenomellinus 90
stevensoni 75, 97
stuckenbergi 92

trevirus 88
turneri 59, 97

* uelleburgi 91, 98

101

FARI

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19:

Re DEEL 117 AFLEVERING 4 1974
Me... MUS. COMP. ZOOL!
| LU LIBRARY

EF e 58-29 | OCT 7 1974
3 4 HARVARD

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| VOOR ENTOMOLOGIE

AR Lp UITGEGEVEN DOOR

| DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

a INHOUD

a D. Hırız Rıs LAMBERS. —On American Aphids, with descriptions of a new genus and
| some new species (Homoptera, Aphididae), pp. 103—155, Figs. 1—13.

% È Tijdschrift voor Entomologie, deel 117, afl. 4 Gepubliceerd 23-VIII-1974

n N n
\
1
4

ON AMERICAN APHIDS, WITH DESCRIPTIONS
OF A NEW GENUS AND SOME NEW SPECIES
(HOMOPTERA, APHIDIDAE)
by

D. HILLE RIS LAMBERS
Bladluisonderzoek TNO, Bennekom, Netherlands

ABSTRACT

The following new taxa are described: Acyrthosiphon pentatrichopus spec. nov, from Rosa
fendleri and Rosa sp., Utah and Colorado, U.S.A.; Acyrthosiphon vandenboschi spec. nov., from
Potentilla glandulosa, California, U.S.A.; Aphis agastachyos spec. nov., from Agastache urticifolia,
Utah and Idaho, U.S.A.; Aphis astragalina spec. nov. from Astragalus sp., Montana, U.S.A.
and Alberta, Canada; Aphis atromaculata spec. nov., allegedly from Ribes sp., Oregon, U.S.A.;
Aphis baccharicola spec. nov., from Baccharis pilularis, California, U.S.A.; Aphis mulini spec.
nov., and Aphis mulinicola spec. nov., from Mulinum sp., Patagonia, Argentine; Masonaphis magna
spec. nov., from Composites, Alberta and British Columbia, Canada; Megourina gen. nov., type-
species Megourina lagacei spec. nov., from Aguilegia formosa, California, U.S.A.; Pentamyzus
falklandicus spec. nov., from Poa flabellata, Falkland Isles; Thripsaphis (Trichocallis) californica
spec. nov., from Carex near C. comosa, California, U.S.A; Thripsaphis (Trichocallis) hybrida spec.
nov., from Carex sp., California, U.S.A; Thripsaphis (Trichocallis) ossiannilssoni pacifica subspec.
nov. from Carex sp., California, U.S.A.; Thripsaphis (Trichocallis) scabra spec. nov. from Carex sp.,
California and Idaho, U.S.A.; Thripsaphis verrucosa nodulosa subspec. nov., from Carex sp., Alaska,
U.S.A., and Labrador, Canada; Thripsaphis verrucosa subverrucosa subspec. nov., from Carex sp.,
California, U.S.A.; Uroleucon sijpkensi spec. nov., from Solidago macrophylla, Quebec, Canada.
The following new synonymies are proposed: Hyperomyzus boerneri Prevost, 1959, and Hyperomyzus
sobrinus F.P.Miiller, 1966, of Neonasonovia zirnitst Hille Ris Lambers, 1952; Dactynotus pieloui
Richards, 1972, of Uroleucon hieracicola (Hille Ris Lambers, 1962) comb. nov, transferred from
Dactynotus; Thripsaphis gelrica Hille Ris Lambers, 1956, of Allaphis caricicola (Mordvilko, 1914);
Callaphis Mordvilko, 1909, type species Callaphis caricicola Mordvilko, 1914, and Allaphis Mord-
vilko, 1921 nom. nov. pro Callaphis Mordvilko, 1909 nec Walker, 1870, of Thripsaphis Gillette,
1917, type species Brachycolus ballii Gillette, 1908; Thripsaphis thripsoides Hille Ris Lambers,
1939, of Allıphis caricis Mordvilko, 1921. New combinations are: Nearctaphis argentinaeradicis
(Gillette & Palmer, 1932), from Aphis; Neonasonovia inflata (Richards, 1962), from Hyperomyzus;
Neonasonovia fronki (Knowlton, 1945), from Amphorophora; Neonasonovia nabali (Oestlund,
1886), from Rhopalosiphum; Neonasonovia nigra (J. McVicar Baker, 1934) (with a redescription of
apterae), from Amphorophora; Neonasonovia thorsteinni (Stroyan, 1960), from Hyperomyzus;
Pentamyzus acaenae (Schouteden, 1904), from Rhopalosiphum; Neobacillaphis striata (Bozhko,
1961), from Thripsaphis; Uroleucon nigrotuberculatum (Olive, 1965), from Dactynotus; Uroleucon
sonchellum (Monell, 1879), from Siphonophora. The apterous viviparous female of Masonaphis
corylina (Davidson, 1914), is described; Aphis mimuli Oestlund, 1887, is redescribed; and the
host alternations of Neonasonovia nabali (Oestlund) and Aphis ribiensis Gillette & Palmer, 1929
are discussed. Thripsaphis (Trichocallis) caricis (Mordvilko, 1921), Thripsaphis (Trichocallis)
foxtonensis Cottier, 1953, and Thripsaphis (Trichocallis) producta (Gillette, 1917) are distinguished
as separate taxa. A key to the American members of Thripsaphis (Trichocallis), and to the sub-
species of Thripsaphis (Trichocallis) verrucosa (Gillette, 1917) is given.

103

104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974
Acyrthosiphon pentatrichopus spec. nov.

Fundatrix.

Colour in life not known. In mounted specimen body rather broadly spindle-shaped.
Tergum faintly sclerotic, not smooth, but wrinkled to irregularly reticulated, the head
pale brownish yellow, the rest not pigmented. Distinct semiglobular marginal tubercles
present on all thoracic segments and on abd. segments II—IV or V, and besides, spinal
tubercles on tergite VIII; the marginal tubercles elevated, on pronotum to 0.030 mm in
diameter, elsewhere smaller. Dorsal hairs not numerous, about 10—14 on each of abd.
tergites I—II, inverted bottle-shaped, on tergites I—III about 0.008 mm long, the 4 hairs
on tergite VIII about 0.016—0.020 mm long. Frontal tubercles rather low, much
diverging, quite smooth, with 4—6 hairs each; median process hardly developed.

Fig. 1. Acyrthosiphon pentatrichopus spec. nov. Fundatrix: a, antenna; b, hind tarsus; c, last rostral
segment; d, cauda; e, siphunculus with probably broken apex. All X 72

Antennae much shorter than body, pale, very gradually darker to the brown processus
terminalis, without noticeably darker tips to the segments; flagellum gradually more
distinctly imbricated from base of segment III to apex; rhinaria on segment III rather
small and inconspicuous, not elevated or protruding, rather far apart up to nearly halfway
the segment; segment III with 22—24 very inconspicuous hairs of 0.006—0.008 mm
long, 1/6—1/4 of basal diameter of the segment; processus terminalis short, with one
hair besides the apical ones. Rostrum just reaching the middle coxae; last segment 0.10
mm long, about 5/7 of second joint of hind tarsi, with 9 hairs besides the 3 subapical
pairs. Legs pigmented like the antennae, apices of tibiae slightly darker, tarsi dark brown;
femora ventrally with many small scales towards apex; first tarsal joints with 5,5,5 or
5,5,4 hairs, the median one half as long as the others; second tarsal joints imbricated,
with about 4 pairs of ventral hairs. Siphunculi cylindrical with wide base, pale without
darker apex, gradually more distinctly imbricated from base to apex, halfway about as
thick as middle portion of hind tibiae; apical part missing. Cauda elongated triangular
with a very faint constriction, pale, with 8 hairs of which only the most basal ones are
acute, the others stunted and partly very short (e.g., 0.013 mm). Subgenital plate with
8 hairs near posterior margin, 2 hairs on anterior half.

Measurements in mm.

No. Length Ant. Ant. segments Siph: Wi CausemeRhint
body III IV V VI on III

1 3.20 2.49 0.75 0.43 0.47 0.21 + 0.41 0.72 0.44 1&3
(from Rosa fendleri, Helper, Utah, U.S.A., 17.V.’58, leg. G. F. Knowlton).

D. Hire Ris LAMBERS: On American aphids 105

Apterous viviparous female.

In mounted specimens body slenderly spindle-shaped. Rather like fundatrix, but tergum
much thinner, indistinctly wrinkled, not reticulated. Spinal and marginal tubercles
distinct, but slightly flatter. Dorsal hairs about 11/,—2 times as long as in fundatrix,
tergite VIII with 5—8 hairs. Frontal tubercles better developed. Antennae much longer
than body, and segments of flagellum with faintly darker tips; segment III with 3—12
rhinaria irregularly spaced in mostly single file over basal 1/3—6/11 part of segment.
Last rostral segment 0.105—0.11 mm, only about 5/9 of second joint of hind tarsi, with
9—11 hairs besides the 3 subapical pairs. Legs much longer; second tarsal joints ventrally
with several more hairs. Siphunculi about 2/7-—1/3 of length of body, about as long as
ant. segment III, with small flange. Cauda less than half as long as siphunculi, more
acute and not or hardly constricted, with 7—9 hairs of which the 3—5 stunted dorso-
apical hairs are longer. Otherwise as in fundatrix.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Siph. Cau.
body III IV V VI on III

1 3629 3.86 0.95 077 0.67 020551205 7&8 1.02 0.43

2 Seb 35971, 0.97 0.74 0.71 02075521210 4&5 0.95 0.41

3 2.99 4.16 0.99 0.78 0.75 0,.207:71218 3&5 0.99 0.43

4 3.36 3.98 1.04 0.79 0.74 0.18 + 1.00 IE 1.01 0.44

(1-4, from Rosa sp., Mud Springs, Colorado, U.S.A., 9.VII.’66, leg. F. C. Hottes &
H. L. G. Stroyan no. U.S.A. 281).

Discussion. The first available specimen, a fundatrix, rather strongly resembles
fundatrices of Acyrthosiphon wasintae (Hottes) as to general shape, length of processus
terminalis, etc., but it differs by the chaetotaxy of the first tarsal joints, the length and
chaetotaxy of the last rostral segment, the caudal hairs, etc. It is very surprising that
Palmer (1952, p. 330, fig. 396) draws the processus terminalis of the male of
A. wasintae as in the fundatrix of our new species, but writes that it is 0.90 mm long.
It is clear that our first A. pentatrichopus is a fundatrix, because an embryo inside has a
comparatively very much longer processus terminalis. Dr. Stroyan kindly allowed me to
add a description of his material, which consisted of apterous viviparae of probably the
third generation.

The species can be separated from all known species of Acyrthosiphon by the com-
bination of its first tarsal chaetotaxy with siphunculi twice or more times as long as the
cauda. In the number of hairs on the first tarsal joints the aphid agrees with Acyrthosiphon
cyparissiae (Koch), but that species has short siphunculi.

Types. Holotype: fundatrix, from Rosa fendleri, Helper, Utah, USA 1g. ASS:
leg. G. F. Knowlton, in the author’s collection. Paratypes: apterous viviparous female,
from Rosa sp., Mud Springs, Colorado, U.S.A., 9.VIL’66, leg. F.C. Hottes & H.L. G.
Stroyan no. U.S.A. 281, in the collection of Dr. Stroyan, Harpenden, Herts.

106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974
Acyrthosiphon vandenboschi spec. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens body elongate spindle-shaped, 3.75
— 4.35 mm long. Tergum membranous, smooth. No marginal or spinal tubercles present.
Dorsal hairs stiff, rather thick, blunt or with faintly incrassate, colourless apices, the
longest spinal and marginal ones on tergite III 0.052 mm long, the shortest 0.026 mm;
tergites I—III each with about 14—20 hairs of which 6—12 are marginal ones; tergite
VII with 10—12 hairs of which the longest are 0.055 mm long. Frontal tubercles large,
diverging, quite smooth, with 8—10 hairs of about 0.050 mm long; median process
hardly developed. Antennae about 1 1/5 times as long as body, pale, with the very tips
of segment III, IV and V, and the part around the rhinaria on VI dark brown;
flagellum, except the very base of segment III, evenly and lightly imbricated; segment I
with 11—14 hairs; segment III without rhinaria, with some 30 rather spreading, stiff
hairs of up to 0.035 mm long, i.e., about 2/3 of basal diameter of segment; processus
terminalis with about 10—19 small hairs besides those at apex. Last rostral segment about
0.22—0.25 mm long, 1 1/2—1 4/7 times as long as second joint of hind tarsi, not
pointed, with about 40 hairs besides the 3 subapical pairs. Legs long, pale, with only —
the apices of the tibiae brown, and the tarsi blackish brown, with rather spiny hairs, the
femora very slightly imbricated; first tarsal joints with 3, 3, 3 spiny hairs of similar shape,
the middle one little shorter than the others; second tarsal joints slightly imbricated, with
numerous ventral hairs. Stigmal pori oval, not or only very faintly reniform, about 0.026
X 0.016 mm. Siphunculi pale with the very apices brown, evenly, basad almost den-
ticulately imbricated from base to apex, nearly cylindrical with widened base, very slender,
at mid-length thinner than middle portion of hind tibiae, very long, about 2/7—3/8 of
length of body, without a trace of reticulation below the well developed, thin, flange.
Cauda pale, not constricted, with almost cylindrical basal half, and evenly tapering distal
half, just not acute, 3/8—2/5 of the siphunculi, with 16—19 subacute to acute hairs.
Subgenital plate with 12—16 hairs along posterior margin and 2—4 on anterior half.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau.
body III IV V VI
i 3.84 4.53 0.98 0.85 0.75 0.18 + 1.46 1.36 052
2 4.04 —— 1.04 0.91 0.75 (O ARI ee 125 0.48
3 4.35 = 1.04 0.96 0.80 OTO 1.43 0.52
4 4.34 4.75 1.12 0.93 0.70 0.17 + 1.48 1.36 0.49
> 4.28 4.52 1.07 0.83 0.75 0.18 71.36 1.38 0.48
6 4.36 4.77 1.08 0.98 0.76 0.200 00659 1.46 0.51

(1—6, from Potentilla glandulosa, Willow Creek (Monterey Co.), California, U.S.A.,
4.1V.’66, leg. R. van den Bosch no. 66-IV-4i and 66-IV-4m).

Alate viviparous female.
Colour in life unknown. Mounted specimens with faintly brownish yellow head,
fuscous around the ocelli, similarly pale mesothorax, minute pale brownish marginal

D. Hire Ris LAMBERS: On American aphids 107

sclerites on abdomen. Shape similar to apterous. viviparous female. Antennae 11/ times
as long as body; basal segments brown, very base of segment III almost colourless but
rest of segment black, the other segments from rather dark brown to blackish, but paler
than segment III; segment III from base to near apex with about 42—60 small to
medium-sized, hardly elevated or protruding, broad-rimmed rhinaria scattered over half
the circumference. Femora brownish yellow, blackish brown dorsally near apex; tibiae
with middle portion brownish yellow, at the very base pale, in between mottled dark
brown, apically black like the small tarsi. Wings with dark, blackish brown, very faintly
bordered veins; stigma pale smoky with dark posterior border. Marginal sclerites each
with some 6—10 closely packed nodules. Siphunculi at apex just wider than near middle,
with pale bases, blackish brown basal 1/2—2/3 part and similar apex, but between the
dark apex and the lower portion mostly paler and more transparent. Cauda elongated
triangular, dusky. Subgenital plate very slightly pigmented. Other characters very much
like those in apterous viviparous female.

n
ita Re

50 120) aap

DARO EE

Fig. 2. Acyrthosiphon vandenboschi spec. nov. Al. viv. fem: a, siphunculus; b, cauda; c, last rostral
segment; d, hind tarsus; f, antenna. Apt. viv. fem.; e, cauda; g, antenna; h, siphunculus. All X 72

108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Siph. Cau.
body III IV V VI on III
1 3:78 4.76 0.99 0.93 0.82 OL 145% 91.8254. 1.34 0.47
2 4.00 5.08 1.02 0.99 0.86 O19 NGL, 153182567147 0.49
3 4.03 5.02 1.03 0.98 0.82 0:20, 1 57 AT So Oey, 0.50
4 3.74 4.62 0.95 0.90 0.77 OM SENS 2 248,853. 01532 0.43
5 3.96 4.86 1.03 0:95 0.83 ONS #55 2527600 0.49
6 3.88 4.89 1.09 0.91 0.79 0.19% 1D OPT 4278509 1532 0.53

(1—6, with apterous viviparous females no. 1—3).

Discussion. This species with its very long and slender siphunculi immediately reminds
one of A. macrosiphum (Wilson) and A. gossypii Mordv., but it is at once distinguished
by the very hairy last rostral segment, and by the absence of secondary rhinaria in apterae
viviparae.

The species is named for my friend and colleague Dr. R. van den Bosch, of the
University of California, Berkeley, who in his hunt for aphid parasites produces a steady
stream of unknown aphids.

Types. Holotype: apterous viviparous female (no. 1 of measurements) from Potentilla
glandulosa, Willow Creek (Highway no. 1, Monterey Co.), California, U.S.A., 4.IV.°66,
leg. R. van den Bosch no. 66-IV-4i. In the authors collection. Paratypes: apterous and
alate viviparous female with collecting data as for holotype, in the collections of the
Biol. Control Dept., Univ. of California, Berkeley, and in the author's collection.

Aphis agastachyos spec. nov.

Apterous viviparous female.

Colour in life dark green (Stroyan). In mounted specimens either quite pale or, in
slightly alatiform apterae, head infuscated, siphunculi gradually darker towards apex,
cauda quite dark like the apices of tibiae and the tarsi. Body about 1.50—2.00 mm long,
broadly oval, with only on the mesothoracic pleura and caudad each siphunculus more
or less pigmented sclerites. Tergum often distinctly reticulated. Dorsal hairs variable in
number, often pleural or duplicated spinal hairs present; hairs stiff, with fine, not
flexuose apices, long, on tergite III spinally up to 0.055 mm long; tergite VIII with
4—8, mostly 6, hairs of similar shape up to 0.070 mm long. Marginal tubercles only
on abdominal tergites I and VII, bluntly conical, about as tall as their basal width (e.g.,
0.020—0.030 mm). Front not much sinuated, with the hairs as long as, but stouter than
hairs more ventrally. Antennae evenly pale or with darker apices, 1/2—5/9 of length
of body; flagellum ventrally markedly imbricated from the very base of segment III, but
segment III dorsally nearly smooth, in true apterae without rhinaria or in specimens
with dark head with 1—5 small rhinaria; processus terminalis 1 1/2—1 4/5 times as long
as base of segment VI, mostly somewhat bent near the primary rhinaria; hairs on segment
III stiff, variable, from blunt with faintly incrassate apices, to very acute, the longest
about 1 1/2—2 times as long as basal diameter of the segment. Rostrum when extended
reaching a little past middle coxae; last segment about 0.105—0.125 mm long, about

D. Hire Ris LAMBERS: On American aphids 109

Fig. 3. Aphis agastachyos spec. nov. Apt. viv. fem.: a, tibia + hind tarsus; c, last rostral segment;
e, head; f, antenna; h, posterior part of abdomen. Al. viv. fem.: b, cauda; d, siphunculus; g, antenna.

All X 72

114 times the very small second joints of hind tarsi, with 4—8 hairs besides the 3 sub-
apical pairs. Legs pale, or, in specimens with pigmented head, with dark apices to the
tibiae; longest ventral femoral hair 0.060—0.075 mm long; dorsal hairs halfway hind
tibia spreading and longer than local diameter of tibia; tarsi small, second joint twice
apical diameter of hind tibia. Siphunculi pale or in alatiforms infuscated towards apex,
rather thick, evenly tapering to near apex, or there just perceptibly constricted, about
1/9—1/7 of length of body, imbricated but apical 1/5—1/4 part at least dorsally nearly
smooth, with a very wide (for an Aphis) flange, about 1 1/3 times as wide as the part
just basad. Cauda thick, more or less bluntly triangular, about as long as its basal width,
darker than siphunculi and 1/2—2/3 of their length, with 7—12 acute, strongly curved
hairs, some of which are placed dorso-apically. Subgenital plate with 10—12 hairs along
posterior margin and 6—10 on anterior half.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau.
body III IV V VI
1 1.54 0.83 0.21 0.12 0.12 OO 0:17 0.15 0.11
2 1897, 0.99 0.28 0.15 0.15 0.11. + 0:18 0.22 0.15
3 1.96 1.03 0.26 0.15 0.16 0:135510;20 0.23 0.15
4 1.80 0.94 0.27 0.14 0.14 0:10 770.18 0.19 0.13
5 1.81 1.04 0.26 0.16 0.16 0.12: 0.21 0.22 0.16
6 1.87 1.02 0.27 0.14 0.15 0.35 0,21 0.22 0.15
7 1.51 0.84 0.20 0.12 0.13 0.11 eh 0.17 0.17 0.15

(All from Agastache urticifolia, U.S.A.; 1—2, 5—6, Cub River Canyon Idaho, 13.VIII.

110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

'65, leg. G.F. Knowlton; 3, Monte Christo, Utah, 7.VII.’66, leg. H.L.G. Stroyan;
4, Wellsville Mountains, Utah, 15.VII.’65, leg. G. F. Knowlton; 7, Logan Canyon,
Utah, 20.VII.’71, leg. G. F. Knowlton).

Alate viviparous female.

Colour. Head, thorax, siphunculi, cauda, antennae, large marginal sclerites dark to
blackish, rest as in apterae. In mounted specimens body 1.60—1.90 mm long, with
blackish head and thorax; large marginal and postsiphuncular sclerites, long bars across
tergites VII and VIII, a broad spinal sclerite on tergite VI and often small, paired spinal
sclerites on tergite V darkish. Antennae dark with the very base of segment III pale,
4/7—3/5 of length of body; segment III with 7—16 very unequally-sized, faintly
protruding rhinaria irregularly arranged along one side; segment IV with 1—4 rhinaria;
segment V sometimes with a secondary rhinarium; processus terminalis at most twice as
long as base of segment VI. Legs faintly brownish to brown with femora distad, and
tibiae apically blackish. Venation normal, greyish brown, not bordered. Siphunculi
blackish, cylindrical to slightly tapering, with the large flange to 11/4, times as wide
as the smooth part just below. Cauda narrower than in apterae, with the “hard portion”
less than half the total length, with up to 14 hairs. Tergite VIII with 4—8 hairs.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau. Rhin. on
body III IV Vi VI III IV

10177 000 9400024 70.157 0:14. 0.141 EOLO MM SAS INI
2001630040 0922710 240 0.13 014 OTO TS 70.142 20.125 1078311832782.
3 184 101 0272 015,016 0110.20 018, 0132 10813831831
4 21.66 20.90 0.26 0.132 :0:137 70.100.187 0-14 70:12 2978311 Aree
>», 1.78 0:93. 0.24 014. 014 0.10+01977097 70.157 9782127 37824
6 41272.) 0:97. . 0.26, (0:15) 0.34 2 .0.1170.20 0.15.72 0.122 7748901082

(1—2, with apt. viv. 1—2; 3, with apt. viv. 3; 4—6, with apt. viv. 4—6).

Oviparous female.

Colour in life not known. In mounted specimens much like apterous viviparous female,
but body more slender. Antennae with 5 or 6 segments. Abdominal tergite VIII with
16—20 hairs. Hind tibiae not or hardly incrassate, with 3—12 pseudosensoria, pigmented
like other tibiae. Siphunculi pale with infuscated apex, flange smaller than in apterae
viviparae. Cauda bluntly conical, with 12—14 hairs. Subgenital plate with middle 1/3
part unpigmented, rather evenly hairy.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau.
body III IV V VI

Il 1.52 0.69 0.14 0.10 0.12 0.10+0.15 0.14 0.13

2 1.36 0.57. 0.17 0.10 0.06+0.16 0.11 0.10

3 1559 0.58 0.17 0.10 0.07 +0.15 0.11 0.11

(1, with apt. viv. fem. no. 3; 2—3, with apt. viv. fem. no. 7).

D. Hire Ris LAMBERS: On American aphids 111

Apterous male.

Colour ín life not known. In mounted specimens head and part of the pro- and
mesothorax as well as marginal sclerites, small sclerites on which the spinal hairs from
mesonotum to abd. tergite VI are placed, small postsiphuncular sclerites, and bars across
tergites VII and VIII dark sclerotic. Antennae dark like head, rather thick, with con-
spicuously imbricated flagellum; rhinaria flat, small, over nearly half the circumference
of segments III—V; hairs on segment III over twice as long as diameter of segment
at its constricted base. Siphunculi dark with the very base pale, slightly tapering,
imbricated, with normal flange. Cauda dark, small, slightly constricted, with 6 hairs.
Remainder more or less as is apt. viv. fem.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. on segment Siph. Cau.
body NV V VI III NT

1 roe O72 WOE 012 O11 007+014 12814. 7&8 4&4 00990109

(1, with apt. viv. fem. no. 7).

Discussion. This species seems to curl the leaves of Agastache urticifolia. Several
samples were received from Dr. G. F. Knowlton, and seven slides from Dr. H. L. G.
Stroyan, who provided his colour notes.

The aphid was first identified as Aphis ramona Swain, known from Salvia mellifera
(syn. Ramona stachyoides) from California. But A. ramona differs by having all the
antennal and dorsal hairs blunt and shorter; by having in apterae viviparae 2, rarely 3
hairs on abdominal tergite VIII; and by the flange of the siphunculi being much nar-
rower, about 1 1/5—1 2/9 times as wide as the smooth part basad in apterae viviparae,
about 1 1/3 times as wide as the thinnest part basad in alatae. Sensoriation of antennae,
rostrum, tarsi, etc., are very much the same, but the cauda has fewer hairs and the
subgenital plate has mostly 2 hairs on anterior half.

Because of the several hairs on the basal half of the last rostral segment, the pale
siphunculi and mostly darker cauda in apterae viviparae, and the sensoriation of the
alatae, Aphis ramona Swain and A. agastachyos spec. nov. strongly resemble the Aphis
oenotherae group of Aphis (A. oenotherae Oestl., A. neomexicana Cock., A. sanborni
Patch, A. varians Patch, etc.). They differ from this group by the conspicuous flange to
the siphunculi.

The occurrence of sexuals in early summer is rare in aphids and exceptional in Aphis
L. The oviparous females contained no eggs, and therefore perhaps later the production
of oviparae might be larger. On the other hand it is possible that eggs are laid very early,
and that may be a reason why this conspicuous species was not found by the specialists
working over Utah and Idaho.

Types. Holotype: apterous viviparous female (measurements no. 1), from Agastache
urticifolia, Cub River Canyon, Idaho, U.S.A., 13.VII.’65, leg. G. F. Knowlton, Paratypes:
apterous and alate viviparae with data as for holotype, and all others mentioned under
the measurements.

112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Aphis astragalina spec. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens body broadly oval, about 2.50 mm
long. Sclerotisation of dorsum variable, from largely membranous with scattered spinal
sclerites, a broken, very irregular spino-pleural bar on tergite IV, a broad, very irregular
and perforated, thick spino-pleural bar across tergite V partly fused with a more regular,
longer bar across tergite VI, to almost completely fused thick spino-pleural bars across
tergites I—IV with the sclerotic bar across tergite IV solidly fused with that across V
and partly with that across VI; marginal sclerites small or large, free; tergites VII and
VIII with free bars, the one on VIII encircling the segment to nearly the subgenital
plate; tergum cephalad siphunculi also on the membranous parts distinctly reticulated
(Montana specimens) or not. Dorsal hairs scarce, spinally on tergite III 0.035—0.052
mm long, the 12 hairs on the sclerite on tergite VIII to 0.065 mm long. Marginal
tubercles on tergites I and VII, bluntly conical to cylindrical with rounded tops, on tergite
VII more or less distinctly complex, with a basal diameter of 0.030—0.035 mm. Head
blackish; front sinuate; frontal hairs long and fine, to 0.060 mm long, dorsal cephalic
hairs from 0.040—0.060 mm long. Antennae about 4/7 length of body, with blackish
basal segments, remainder pale with brownish apices to segments (III), IV and V; VI
brown; segment III without scaliness on one side, along the other side markedly im-

Fig. 4. Aphis astragalina spec. nov. Apt. viv. fem.: b, antenna; c, abdomen; d, last rostral segment;
e, tip tibia + hind tarsus. Al. viv. fem.: a, antenna; f, siphunculus; g, cauda. All X 72

D. HiLLE Ris LAMBERS : On American aphids 113

bricated; processus terminalis short, 11/ times base of segment VI; hairs on segment III
rather numerous (8—12), spreading to erect, up to 0.052 mm long, to twice diameter
of segment at its constricted base; segment VI on base with 2—3 hairs of 0.025—0.035
mm, but besides with 1—3 short (0.0085 mm) spiny hairs similar to the 3—4 occurring
on basal 2/3 part of the processus terminalis which besides has 4 on the top. Rostrum
reaching to middle coxae; last segment rather blunt, 0.12—0.13 mm long, 4/5—1 times
as long as second joint of hind tarsi, with 2 hairs besides the 3 subapical pairs. Legs
rather short and thick; femora blackish brown with pale base; tibiae pale with the very
base infuscated and distal 1/4—1/3 blackish brown, conspicuously tapering from near
base to apex; femoral hairs fine, longest ventral one up to 0.070 mm; tibial hairs dorsally
fine and erect, to 0.065 mm long, halfway tibiae just shorter than local diameter of tibia,
but ventral hairs more spiny and on basal part some only 0.022 mm long; first tarsal
joints with 3, 3, 2 hairs; second joints with distal dorsal pair not longer than middle dorsal
pair of fore and mid legs. Siphunculi black, thick, tapering, 1/9—1/8 of length of body,
strongly but rather bluntly imbricated, with just indicated flange. Cauda black, thick,
fingertip-shaped with a faint constriction, just shorter than siphunculi, with 17 normal,
long, curved hairs. Subgenital plate with 10—12 long hairs on anterior half.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau.
body III IV V VI

1 2.40 1.28 0.36 0.23 0.20 Oss 0:19 0.28 0.26
2 2:52 1.44 0.42 0.28 0.24 0.14 + 0.21 0.30 0.27

(1, from white flowered Leguminosa, 4 mi. E. of Elliston, Montana, U.S.A., 20.V11.’66,
leg. F.G.A.—A.P.G. no. G66—VII—20f; 2, from Astragalus, 2 mi. E. of Conmore,
Alberta, Canada, 17.VII.'66, leg. F.G.A.—A.P.G. no. F66—7—17h).

Alate viviparous female.

Colour in life not known. In mounted specimens head and thorax black sclerotic,
abdomen on tergites I—IV with scattered, transverse spinal and pleural sclerites, and
with very large marginal sclerites; on tergite V with an irregular transverse bar, on
tergites VI—VIII with bars as in apterae. Hairs shaped as in apterae, but tergite VIII
with 8 hairs though in view of the irregular arrangement other numbers probably occur.
Antennae with black basal segments, flagellum with the very base pale, rest brown, very
gradually blackish brown towards apex; segment III with the rhinaria very unequal in
size, more or less in.a line; the rhinaria evenly spaced over length of segment IV; hairs
on segment III shorter, to 0.030 mm, but on segment VI as in apterae. Wings probably
with normal venation (damaged), veins brownish grey, not bordered. Siphunculi thinner
and less tapering. Cauda with triangular base and cylindrical or even very elongate oval
distal half, very blunt, with 14 hairs. Other characters as in apterae.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. on Siph. Cau.
body III IV V VI III IV

JAR 2: AS TLC 70:44, 4034.030,1 (0154022). 1.821, 31822 0:23, 0:20
(1, with aptera no. 1).

114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Discussion. Few Aphis spp. have been described from North America after they were
cleared, and therefore it is not possible to discover from published descriptions what kind
of sclerotisation occurs in apterae viviparae of Nearctic Aphis. There appears to be a
small group of Nearctic species that are comparable to the craccivora group of Aphis,
originally Palaearctic. The Nearctic ones I saw, differ by three characters: the presence of
a sclerotic band encircling abd. segment VIII to near the subgenital plate, and bearing
some 3—12 hairs, marginal tubercles on abdomen with a rather wide sclerotic basal part,
and a rather high number of spiny hairs on the processus terminalis and often the basal
part of antennal segment VI. In this sense also Aphis lupini Gill. & Palmer, according
to a specimen identified by Prof. Palmer that I remounted, without a sclerotic dorsum,
comes near this group. None of the species so far known to me has such long antennal
hairs as A. astragalina spec. nov., or so many caudal hairs.

Types. Holotype: Apterous viviparous female (no. 1 of measurements), from white
flowered Leguminosa, 4 mi. E. of Elliston, Montana, U.S.A., 20.VII.'66, leg. F.G.A.
—A.P.G. no. G 66—VII—20 f. Paratypes: one alate vivipara with collecting data as for
holotype, and one aptera vivipara from Astragalus sp., 2 mi. E. of Conmore, Alberta,
Canada, 17.VII.’66, leg. F.G.A.—A.P.G. no. F66—VII—17h.

Aphis atromaculata spec. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens body about 2.10—2.40 mm long,
very broadly oval. Head and thorax black sclerotic; over abd. tergites I—VII thick, nearly
solid, black sclerotic bars, often more or less completely fused on tergites II—VI, the
bars on tergites I, VII and VIII always free; sclerotisation laterally reaching to near the
stigmata, and rather distinctly reticulated. Tall (to 0.045 mm), large (basal diameter
0.035 mm), often constricted, on the basal portion sclerotic, marginal tubercles on abd.
tergites I and VII, in one out of 11 specimens with a smaller one on tergite II. Dorsal
hairs scarce, rather stiff, pointed, on abd. tergites I—II spinally about 0.035—0.045 mm
long, the 3—6 hairs on tergite VIII with fine apices, to 0.070 mm long. Front slightly
sinuated, antennae about 4/7—2/3 of length of body; basal segments black, segments
III and IV pale with brown apices, V and VI dark brown; segment III from base
imbricated by smallish scales, without rhinaria, processus terminalis 1 1/2—1 2/3 times
base of last segment, with 1—2 small spines like the 4 on apex; hairs on flagellum rather
acute and spreading, longest ones on segment III to 0.035—0.040 mm and slightly
longer than diameter of segment at its slightly constricted base. Rostrum according to
stylet length reaching to hind coxae; last segment about 0.14—0.15 mm long, normal,
with 2 hairs besides the 3 pairs near apex. Mid and hind femora blackish with only
the very bases pale, fore femora paler; all femoral hairs except some dorso-apical ones
long and fine, largest ventral hair 0.050—0.060 mm long; tibiae brownish yellow with
the very apex, and distal 1/4—1/3 part blackish; dorsal tibial hairs, excepting 1—4 blunt
ones near base, long and fine, spreading, on basal half to 0.055 mm long, not quite as
long as local diameter of tibia; first tarsal joints with 3, 3, 3, or (perhaps) sometimes
3, 3, 2 hairs; second joints about 0.10 mm long, only 11/ times as long as apical diameter
of tibiae. Siphunculi quite black, more or less cylindrical with widened base, 1/7—1/6
of length of body, evenly imbricated, with very small flange. Tergite VIII with a long
blackish sclerotic bar reaching nearly to the subgenital plate. Cauda black, rather thick,

D. Hire Ris LAMBERS: On American aphids 115

Fig. 5. Aphis atromaculata spec. nov. Apt. viv. fem.: a, tip tibia + hind tarsus; d, last rostral segment;
e, antenna; f, posterior part of abdomen; h, head. Al. viv. fem: b, antenna; c, siphunculus; g, cauda.
AIR 72

fingertip-shaped, perhaps sometimes constricted, 2/5—3/5 of length of siphunculi, with
7—9 almost hooked hairs. Subgenital plate with 5—9 hairs on disc and anterior half.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau.
body III IV V VI

1 215 122 0.32 0.21 0.19 0.14 + 0.21 0.30 0.18
2 2.14 1.28’ ‘0735 0.21 0.21 0.14 + 0.21 0.38 0.18
3 2.28 1.23 0.36 022 0.22 O15 > 0:19 0.36 0.19
4 236 1.54 0.45 0.30 0.24 0.15 + 0.24 0.33 0.18

(IL “Ribes or?”, Zigzag, Oregon, U.S.A., 28.VI.’57, leg. G. F. Knowlton & D. W.
Davis).

Alate viviparous female.

Colour in life not known. In mounted specimens body 2.05—2.45 mm long. Head
and thorax black sclerotic; abdomen on all tergites with completely free, blackish, spino-
pleural transverse bars sometimes fused with intersegmental sclerites but not with the

116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

conspicuous marginal sclerites. Antennae black with only the very base of segment III
brown; segment III with 22—29 medium-sized, nearly flat, not bulging, scattered
rhinaria; segment IV with 0—4 inconspicuous rhinaria in file; segment V only on one
antenna (with 4 rhinaria on IV) with a secondary rhinarium; processus terminalis as in
apterae 1 1/2—1 2/3 times as long as base of segment VI. Wings with normal venation,
veins rather dark brown, stigma about as dark as dorsal abdominal sclerites, darker than
subcosta. Tibiae with more black at base and apex, in old specimens nearly wholly dark;
dorsal tibial hairs more adpressed than in apterae, shorter, and a larger basal portion
with blunt hairs. Siphunculi rather shorter than in apterae. Cauda distinctly constricted,
with 8—12 hairs. Other characters rather as in apterae.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. on Siph. Cau.
body III IV V VI "NAT IV

1 2.30 ve 1:40) 0.37. 02065 40/2208 LO LG 0.23" 225,626 77.8200 274018

2 231 143 0.38 0:24, 40.23. 0.160.260 23% 27 3&0 “O27 016

2, ek 1.57% OS 0:23) 10220 0.157025 2216 22065 2520288047

A 22721] 1.34. 20:39, 2022770227 0:15-0:25 24829 1 &1 2027410218

(1—4, with the preceding morph).

Discussion. This Aphis belongs morphologically to the long-haired section of the
Aphis craccivora Koch group, which has very few members in Europe, all on Legu-
minosae. In America, native long-haired Aphis with extensive black abdominal sclero-
tisation so far are known from only one Leguminosa, Astragalus spp. Aphis cercocarpi
Gill. & Palmer from Cercocarpus spp. seems to be the only earlier described species.
It can easily be recognized by the absence of rhinaria on ant. segment IV in alatae, by
the shape of its cauda, and the pigmentation of the antennae. Apterae have scattered
areas of sclerotisation on the abdomen, as specimens remounted from slides received from
Professor Palmer showed. The present species can easily be distinguished from astragalina
by the much larger number of rhinaria in alatae, and the shorter tarsi in all known
morphs. Similar Aphis species from Spiraea, as yet undescribed, has (or have) also longer
tarsi. The South American A. mulini, described on p. 121 of this paper, differs by
possessing marginal tubercles also on abd. segments II—IV, while in the other American
species they are only present on segments I and VII.

Unfortunately the host plant is not known, but the species can so easily be recognized
that I venture to describe it from the sample. Ribes is the most unlikely host plant.

Types. Holotype: apterous viviparous female (no. 1 of measurements), from “Ribes
or 2”, Zigzag, Oregon, U.S.A., 28.VI.’57, leg. G. F. Knowlton & D. W. Davis no. 367.
Paratypes: the measured apterae viviparae and 7 alatae viviparae.

Aphis baccharicola spec. nov.
Apterous viviparous female.

Colour in life not known. In mounted specimens head and parts of thorax quite dark
to blackish; abdomen with rather short spinal to spino-pleural dark sclerotic bars across

D. Hure Ris LAMBERS: On American aphids 117

tergites VII and VIII, and often a short, thicker spinal sclerite on tergite VI; small
marginal sclerites and large post-siphuncular sclerites on tergites I—VI but these paler
than the blackish intersegmental sclerites; sometimes small fragments of spinal sclerites
on more anterior tergites. Tergum more or less distinctly reticulated. Dorsal hairs sparse,
fine, with wavy apices, the spinal ones on abdominal tergite III about 0.050—0.060 mm
long, the 2—4 hairs on tergite VIII to 0.070 mm long. Marginal tubercles on abdomen
only on tergites I and VII, large, less than half as long as their basal width (e.g.,
0.045—0.060 mm). Front sinuated, the frontal and dorsal hairs in length similar to the
ventral ones. Antennae with basal segments slightly paler than head, flagellum pale with
dark apex; length of antennae 5/8—2/3 of length of body; segment III without rhinaria,
rather sharply imbricated from near base; processus terminalis 1 8/11—2 3/10 times base
of segment VI, usually with 2—3 spines besides those at apex; hairs on segment III
conspicuously erect, fine, the longest ones just over twice as long as diameter of the
segment at its only slightly constricted base. Rostrum when extended reaching to hind
coxae; last segment rather slender, 0.11—0.12 mm long, subequal to second joints of
hind tarsi, with 2 long hairs besides the 3 subapical pairs. Femora blackish brown with
shorter or longer pale basal part; longest ventral femoral hair about 0.060 mm; tibiae
with the very base and distal 1/5 part dark brown, the rest yellowish to brownish yellow;
dorsal hairs on hind tibiae spreading, all long and fine, halfway tibia about 0.060 mm
long, 11/2—1 3/5 times as long as local diameter of tibia; hairs on ventral side of
tibia more variable, all shorter, and mostly more spiny. Siphunculi black, rather thick,
evenly densely imbricated, tapering to the middle with distal half cylindrical or tapering
to the top, with the apex often faintly attenuated, in the middle about 1 3/8—1 1/2
times as thick as halfway diameter of hind tibiae, about 1/6—2/11 of length of body,
rather often with up to 2 hairs of the type of the dorsal body hairs but shorter; flange
small, about 11/10 times as wide as the part just basad. Cauda as dark as siphunculi,
very strongly tapering on basal 1/3 part, less tapering on distal part, quite blunt,
1/2—2/3 of length of siphunculi, with 8—9 very strongly curved hairs, one of which
is dorso-apical. Subgenital plate with 2 hairs on anterior half.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau.
body III IV V VI
1 1.58 1.09 0.31 0.17 0.15 Os E10: 25 027 0.15
2 1072 0.98 0.28 0.15 0.14 OA1=270.19 0.31 0.15
3 1.45 0.96 0.22 0.15 0.14 OR O22: 0.23 0.14
4 1.63 0.96 0.28 0.14 0.14 0.10°+ 0.19 0.27 0.11
5 1.45 0.98 0.26 0.14 0.14 0.10 + 0.23 0.25 0.14
6 1.63 0.98 0.26 0.16 0.14 OMO EE 021 0.27 0.14
7 1.60 1.00 0.27 0.15 0.14 0.10 + 0.23 0.26 0.13

(1, from Baccharis pilularis, Patrick Point State Park (Humboldt Co.), California,
U.S.A, 19.VIII. 66, leg. C. Lagace and D. Calvert no. G 66—VII—19 g; 2—7, from
Baccharis sp., Clam Beach, Highway 101 (Humboldt Co.), California, U.S.A., 19.VIII.
‘66, leg. C. Lagace, no. L 66—VIII—19 e).

118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Alate viviparous female.

Colour in life not known. In mounted specimens head and thorax black, ahdomen with
similar sclerotic pattern as in apterae, but spinal sclerotisation much more extensive,
though marginal sclerites still very little developed. Antennae blackish with base of
segment III brown to pale brown, about 3/5—2/3 of length of body; segment III with
about 14—22 rather equally sized, slightly protruding, rather small rhinaria along one
side not in line; segment IV without rhinaria, processus terminalis to 2) times base
segment VI. Legs more slender and darker than in apterae. Venation of wings normal,
with second fork of media sometimes rather near apex; veins thick and brownish, not
bordered, siphunculi less thick than in apterae, with or without hairs. Cauda more evenly
tapering. Tergite VIII with 3—6 hairs. Other characters as in apterae viviparae.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau. Rhin.
body III IV V VI on III

1 ail 7 9.33 0.18 0.16 OS 40,25 0.26 0.14 19 & 22
2 1.95 aad OY 0.34 0.19 0.16 02025 0.30 0.15 16 & 20
3 1256 1.14 0.31 0.17 0.16 OC NOED 0.23 0.12 14&17

(1—3, with aptera no. 1; 3, intermediate with wing pads).

Fig. 6. Aphis baccharicola spec. nov. Apt. viv. fem.: a, siphunculus; c, last rostral segment; e,
posterior part of abdomen; f, antenna. Al. viv. fem.: b, siphunculus; d, cauda; g, antenna. All X 72

Discussion. Every detail of the morphology of this species suggests that it belongs
to the Aphis helianthi Monell group of the genus. Also A. ceanothi Clarke belongs to
this group, which is characterized by a comparatively long third antennal segment with
numerous irregularly arranged rhinaria in alatae, no rhinaria on segment IV, a rather
short processus terminalis, moderately short antennal hairs; long and fine dorsal body
hairs; thick, dark siphunculi and an almost equally dark cauda with 7—10 hairs; 2 hairs
on the anterior part of the subgenital plate; on abdomen marginal tubercles only on
segment I and VII; 2 hairs besides the 3 subapical pairs on the last rostral segment.
Within this group Aphis baccharicola can immediately be recognized by its very long and
fine antennal hairs, large marginal tubercles, and the not infrequent occurrence of long
hairs on the siphunculi.

The new species seems to be widely spread on Baccharis pilularis, together with its

D. Hire Ris LAMBERS : On American aphids 119

subspec. consanguinea a common shrub in the California coastal area

Types. Holotype: apterous viviparous female (without hairs on siphunculi, measure-
ments no. 1), from Baccharis pilularis, Patrick Point State Park (Humboldt Co.),
California, U.S.A., 19.VIII.’66, leg. C. Lagace & D. Calvert no. G 66—VIII—19 g.
Paratypes: alate viviparae (with hairs on siphunculi) with collecting data as for holotype;
apterae viviparae (some with hair on siphunculi), from Baccharis sp., Clam Beach, High-
way 101 (Humboldt Co.), California, U.S.A., 19.VIII.’66, leg. C. Lagace no. L 66—VIII
UNE

Aphis mimuli Oestlund, 1887

Apterous viviparous female.

Colour in life not known. In mounted specimens quite pale with only the apices of
antennae and tibiae brown and the cauda rather dark. Body about 1.55—1.75 mm long,
rather broadly oval, without sclerotic pigmented spots. Reticulation of tergum faintly
visible. Dorsal hairs sparse, stiff, blunt, on abdominal tergite III spinally about 0.015
—0.026 mm long, caudad much longer, the two hairs on tergite VIII very acute and up
to 0.060 mm long. Marginal tubercles only on abdominal tergites I and VII, conical,
about half as long as their basal width of about 0.026 mm. Front rather sinuated, with
the hairs 0.025—0.035 mm long. Antennae pale with the apex from basal one-third part
of segment V more or less dark brown, 4/9—5/9 of length of body; flagellum on basal
1/3—1/2 of segment III ventrally with some small dispersed scaly imbrications, more
distally strongly but not densely imbricated, without secondary rhinaria; processus ter-
minalis 1 4/5—2 1/9 times as long as base of segment VI; segment III with about 5—8
stiff, blunt hairs, the longest of which is just shorter than basal diameter of the segment.
Rostrum nearly reaching hind coxae; last segment about 0.13—0.14 mm long, with about
6—9 hairs besides the 3 subapical pairs. Legs pale with apices of tibiae brown; femora
hairy, longest ventral hair about 0.025 mm long; hind tibiae with rather erect, stiff hairs
with mostly spear-shaped apices, halfway the tibiae about 0.019—0.020 mm long, just
shorter than local diameter of the tibia. Siphunculi pale with sometimes the apex faintly
dusky, rather thick, tapering to distal one-fourth part or to apex, just attenuated, 2/15
—2/11 of length of body, on basal half almost smooth, on distal half superficially im-
bricated. Cauda dark, slightly constricted, with fingertip-shaped distal part, 3/5—2/3
of the siphunculi, with some 12—16 curved, normal hairs. Subgenital plate with some
16—20 hairs along posterior margin, 6—10 long hairs on anterior half.

Fig. 7. Aphis mimuli Oestlund. Apt. viv. fem.: a, tip tibia + hind tarsus; b, siphunculus; c, cauda;
h, antenna. Al. viv. fem.: d, siphunculus; e, cauda; f, last rostral segment; g, antenna. All X 72

120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau.
body III IV V VI

L 1465 0.84 ORT 0.14 0.14 0.105,08 0.22 0.15

2 197 0.87 Oi, 0.16 0.14 0:10%=#30:19 0.28 0.16

3 1.66 0.81 0.17 0415 0.12 0.10 + 0.18 0.25 0.14

4 175 0.84 0.17 0.14 0.13 0.09 2019 0.23 ?

(1—3, from Mimulus aurantiacus, Union Landing (Mendocino Co.), California, U.S.A.,
18.VIII.’66, leg. D. Calvert no. C 66—VIII—18 b; 4, from Mimulus guttatus, Plaskett
Creek (Monterey Co.), California, U.S.A., 5.IV.’66, leg. R. van den Bosch no. 66—IV

Alate viviparous female.

Coulour in life not known. In mounted specimens head and thorax black; on the
abdomen rather large marginal and postsiphuncular sclerites, a thin spinal bar on tergite
VIII, a thicker spinal bar on tergite VII, a very thick rectangular spinal bar on tergite VI,
a complete to rudimentary spinal bar on tergite V and some small irregular sclerites on
more anterior tergites dark, but paler than the conspicuous intersegmental sclerites.
Antennae blackish with thick, imbricated flagellum, 5/9—3/5 of length of body; segment
III up to twice as thick as its basal width, with 17—21 rather equally sized, broad-
rimmed rhinaria over most of its circumference; segment IV with 7—11 rhinaria;
segment V with 0—4 secondary rhinaria, and even basal part of segment VI sometimes
with a secondary rhinarium; processus terminalis 2—2 1/5 times as long as base of
segment VI. Legs dark, with the basal part of the femora and the larger part of the tibiae
only in immature specimens pale, but then still the very base of the tibiae dark. Venation
of wings normal; veins dark, rather heavy but not distinctly bordered; stigma blackish
grey. Siphunculi blackish, nearly cylindrical, imbricated from near base, shorter than in
apterae, with the distinct flange nearly 1 1/5 times as wide as the part just below. Cauda
rather slender, dark with paler tip, with only 9—11 hairs. Anterior part of subgenital
plate with (3)4—8 hairs. Other characters as in apterae.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau. Rhinaria on segments
body TRIS INVE SV VI III IV ME AVI
1 1.87) 1.04 0.27 30.17. (0.116; (0.11-F0:23 10:22) 0.15 (18821 TOEG 3844080
2 1:89. 1.00% 0:25 00:17M0:16 0100210 190130178177 SO RETZ MELO
3.065, 11-00 0:26 40:15, 7015) 01102-02309) POET OECD ONMS SIND SES MAESEN
4 1,74 1.09. 0:29 10:17 0517, 0:11 20-237 10:21 770.14 17.821721 08:3 MIE ROSEN
5). 1:63. 0:92: 0:23 10.15 0:13 0.097.027 TOU OL 1882S S320 ROO

(1—5, with apterae 1—3).

Discussion. This is a typical member of the Aphis oenotherae group of species, which
so far was only known to occur on Ribes spp. and on Onagraceae (Oenothera, Epilobium,
Chamaenerium). IÉ one sample from Mimulus were available, it would not have been

D. Hire Ris LAMBERS: On American aphids 121

described here, but there are two, from different species of Mimulus. If both apterous
and alate viviparae are available, samples can be identified. Apterae look like Aphis
oenotherae Oestl. or A. neomexicana Cock., but alatae have too many rhinaria. Palmer
(1952) mentions 11—22 rhinaria in alate A. oenotherae but the high numbers almost
certainly relate to Aphis varians Patch, which goes from Ribes to Chamaenerium angusti-
folium, also in Colorado. The latter species has a longer last rostral segment, more hairs
on the antennae, and as a typical character 4—8 hairs on abdominal tergite VIII, besides
considerably longer antennal hairs. Therefore the chaetotaxy distinguishes Aphis mimuli
from A. varians Patch, the number of rhinaria separates it from the species described as
A. sanborni Patch, A. ribiensis Gill. & Palmer, A. ribigillettei Allen & Knowlton and
A. oenotherae Oestl.

It seemed desirable to publish a redescription of the species which was very in-
sufficiently described in 1887 and briefly mentioned in Oestlund (1922). I did not find
any further descriptive notes.

Aphis mulini spec. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens body broadly oval, about 1.75—2.00
mm long. Sclerotisation of body variable, consisting of thick, very variable, conspicuously
reticulated, blackish spino-pleural transverse bars on body from metanotum to abd.
tergite VIII; in very sclerotized specimens these bars tend to fuse on tergites I—V, maybe
only two of them; in little sclerotized specimens the bars show perforations and a median
split on tergites III, and as a limit are reduced to paired, very irregular and fragmented
spino-pleural sclerites; marginal sclerites on abd. tergites II—IV absent, but traces present
on tergite V; spino-pleural bar on tergite VIII short, thin or absent; pleural inter-
segmental sclerites on abdomen darker than other sclerotic parts, sharply bordered, and
looking like groups of black yeast cells. Dorsal hairs on abdomen on tergites I—IV both
pleurally and spinally present, but on nearly all of these tergites one or more spinal or
pleural hairs absent; longest spinal hairs on tergite III to 0.039 mm long, acute or with
a fine almost invisible apex; the 2—4 hairs on tergite VIII up to 0.060 mm long.
Marginal tubercles darkish, cylindrical with rounded tops to bluntly conical, besides on
tergites I and VII (basal diameter 0.030—0.040 mm) also regularly on tergites II—IV
and VI on which they are about half that wide. Head blackish; front sinuate; frontal
hairs fine, to 0.048 mm long; dorsal cephalic hairs at least as long. Antennae about 3/5
of length of body, with blackish basal segments and pale flagellum, apparently without
darker apices to segments III—V; segment III without rhinaria, ventrally imbricated,
dorsally apparently smooth, though other segments on both sides imbricated, with fine,
erect hairs of up to 0.035 mm, about 1 1/3 times as long as diameter of segment at its
somewhat constricted base; segment VI about as long as III, with short processus ter-
minalis. Rostrum not quite reaching hind coxae; last segment about 0.115—0.125 mm
long, subequal in length to 2nd joint of hind tarsi, with 2 hairs besides the 3 subapical
pairs. Legs pale with dark apices especially the hind femora, and tibiae; longest hair on
trochanter 11/5 times as long as suture between trochanter and femur; dorsal hairs
on tibiae long and fine, inclined, on basal half to 0.055 mm long, distinctly longer than
the more spiny ventral hairs, and longer than local diameter of tibiae; first tarsal joints
with 3, 3, 2 hairs; second joint of hind tarsi to 3 times as long as apical diameter of

122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

hind tibiae. Siphunculi evenly black, with dense, rather long, subacute imbrications over
whole length, about 1/6 of length of body, rather evenly tapering from base to the small
flange. Cauda blackish, fingertip-shaped, about 2/3 of length of siphunculi, with 10—14
long, curved hairs with fine apices. Subgenital plate with 10—12 hairs on posterior part,
and 2—3 on anterior half.

Measurement in mm.

No. Length Ant. Ant. segments Cau. Siph.
body III IV V VI

1 1.78 1.08 0.28 0.19 0.19 ORE 0.17 0.28 0.20

2 1.77 all 0.26 0.20 0.18 012021 0.28 0.20

3 1.94 1.10 0.28 0.19 0.17 O12 0:20 0:32 0.22

4 1.92 1.15 0.29 0.20 0.18 0.12 7 0:20 0.31 0.23

(1—4, from Mulinum sp., Bariloche, Neuquen, Argentine, 15.1.'64, leg. D. C. Lloyd).

Discussion. The four specimens were in very good condition in an excellent balsam slide
received from Dr. R. C. Dickson, Riverside. I received no reply to a letter with a request
for further information about the material. Because the slide contained two very similar
Aphis spp., it had to be remounted. During the process specimens came in contact with
a rubbery glue used for fixing labels, and they were badly damaged. Fortunately the
measurements and description were made before remounting.

Because abdominal tergite VIII has only a very short transverse sclerotic bar, and only
2 hairs, this species is not affiliated to North American Aph?s with a markedly sclerotized
dorsum in apterae viviparae. It has all the characters of the European craccivora, or rather
medicaginis group, for abd. tergites II—IV show consistently marginal tubercles. In that
group only one species is known with long antennal, tibial and dorsal body-hairs: Aphis
comosa Börner. But that species has only 6—9 caudal hairs, and a solid sclerotic dorsal
shield that does not tend to be divided into separate bars, but is merely becoming reduced
on its anterior part in specimens that are starved. All other species with rather long
siphunculi have much shorter hairs.

Fig. 8. Aphis mulinicola spec. nov. Apt. viv. fem.: a, posterior part of abdomen; e, antenna.
Aphis mulini spec. nov. Apt. viv. fem.: b, siphunculus; c, last rostral segment; d, head; f, antenna;
g, cauda. All X 72

D. Hire Ris LAMBERS: On American aphids 123

Types. Holotype: apterous viviparous female (measurements no. 1), from Mulinum
sp., Bariloche (Patagonia), Argentine, 15.1.°64, leg. D. C. Lloyd. Paratypes: apterae
viviparae with collecting data as for holotype.

Aphis mulinicola spec. nov.

Apterous viviparous female.

Colour unknown. Strikingly like the preceding species. Differing in the following
characters: dorsal sclerotisation much more extensive, consisting of a large spino-pleural,
intersegmentally hardly perforated patch which on tergite V is linked to the marginal
sclerites; marginal sclerites present, and bearing the marginal tubercles; tergite VIII with
a rather long sclerotic transverse bar bearing 2—4 hairs. Marginal tubercles lower, but
similar as to size and arrangement. Antennae with segments I—V similar in length to
those in A. mulini, but processus terminalis not longer than basal part of segment VI. Last
rostral segment 0.14 mm long, just longer than second joint of hind tarsi. Siphunculi only
1/10 of length of body, almost without a flange, thick, slightly curved outwards. Cauda
thick, as long as or longer than siphunculi, with 8—10 normal, curved hairs. In all other
respects like Aphis mulini.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau.
body III IV V VI

1 1.81 0.99 0.26 0.19 0.16 0413, 270.12 0.17 0.20
2 1.84 1.06 0.28 0.20 0.18 0.130213 017 O7

(1—2, from Mulinum sp., Bariloche, Neugen, Argentine, 15.164, leg’ DY ENT loya)r

Discussion. The two available specimens were in a slide with the preceding species,
but survived the remounting process slightly better. They were first mistaken for fun-
datrices of Aphis mulini spec. nov. I then compared a great number of fundatrices with
apterae of later generations of similar members of the genus Aphis. It became evident,
that, if they were fundatrices, there probably would have been either 5 antennal segments,
or also ant. segments III—V should have been distinctly shorter than they were in
Aphis mulini spec. nov. The date of collecting 15.164 at Bariloche, which is about
41° S. of the equator, would be comparable with 15 July at New York on the Northern
Hemisphere, at which date one could expect fundatrices only as far north as Greenland,
or near a glacier.

Also this species belongs in the craccivora group because of its dorsal sclerotisation,
but species with such long hairs, marginal tubercles on tergites II—IV, and siphunculi
not longer than the cauda, were not previously found in that section of Aphis L.

Types. Holotype: apterous viviparous female (measurements no. 2), from Malinum
sp, Bariloche (Patagonia), Argentine, 15.164, leg. D. C. Lloyd. Paratype: apterous
viviparous female with collecting data as for holotype.

124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974
Aphis ribiensis Gillette & Palmer, 1929
Syn.: Aphis nasturtii Palmer, 1936, 1952 nec Kaltenbach, 1843.

On 17-IX-1956 the late Dr. F. C. Hottes and I found at Carson’s Hole near Grand
Junction, Colo., U.S.A., a small Aphis on stems and leaves of Veronica americana var.
crassata. In a plastic bag with some water the larvae developed into alate females, males,
and one ovipara. This suggested that the Aphis, which in life strongly resembled Aphts
nasturtii Kltb., had host alternation.

The mounted aphids fitted the description of Palmer (1936, 1952) of Aphis nasturtu,
but they differed from the real nasturtii KItb. by having much shorter siphunculi and
processus terminalis. Palmer described her A. nasturti from various Rorippa species, some
of which are also hosts of Aphis nasturtii Kltb.

Through the kindness of Dr. Fronk it became possible to examine most of the samples
to which Palmer (1952) refers. Undoubtedly the Aphis from Veronica is the same as
Palmer’s A. nasturtii. But in her slide from Rorippa, Ft. Collins, Colo., 21-VIII-1918, leg.
L.C.B(ragg) there is also an aptera and an alate of the real A. nasturtii Kltb. And in
one from Twin Lake, Pingree Park, Colo., from Rorippa obtusa, 15-VIII-1932, leg.
M.A.P(almer) there is also one aptera of A. nasturtii Kltb. The latter aphid is described
by Palmer (1952) as A. abbreviata Patch from Rhamnus cathartica, Ft. Collins, a synonym
of A. nasturtii Kltb. and the identification appears to be correct.

Palmer (1952) wrote under Aphis ribiensis Gillette & Palmer: “this species seems to
show no distinguishing characters from Aphis nasturtii; however, since no host transfers
have been done the two species will be held distinct for the present”. To European aphid
workers the identity looks improbable, but this is not the case when one knows what
Palmer meant by Aphis nasturtii. Type material of A. ribiensis was received, with the
kind permission to remount, from the U.S. National Museum and from the Colorado
A. & M. College, Ft. Collins, Colo. I have no doubt that Aphis ribiensis Gill. & Palmer
is the aphid from Rorippa spp. and Veronica americana. Apterae, alatae and oviparae
fully agree. Typical are the protected spiracles on the abdomen, of a type often found
in aphids living in very wet surroundings.

A. ribiensis is the only American Aphis from Ribes with only two accessory hairs
on the last rostral segment as correctly figured by Palmer (1936, 1952). However,
specimens identified by Prof. Palmer as Aphis ribiensis, from Ribes aureum, Nibley,
Utah, leg. G. F. Knowlton, show 5—7 accessory hairs on the last rostral segment which
is 0.12 mm long. The latter specimens agree best with Aphzs ribigillettei Knowlton &
Allen which is virtually indistinguishable from Aphts oenotherae Oestlund.

I should like to point out the similarity in structure and life cycle between Aphis
ribiensis and the European Aphis triglochinis Theob., both with only two accessory hairs
on the last rostral segment, and having host alternation between Ribes and aquatic or
semiaquatic plants.

Masonaphis (Ericobium) corylina (Davidson, 1914) ?

Apterous viviparous female.
Colour in life not known. In mounted specimens body broadly spindle-shaped, about
2.50—2.70 mm long. Tergum not distinctly sclerotic, locally with fine, long, irregular,

D. Hire Ris LAMBERS : On American aphids 125

transverse ridges, especially on the anterior abd. tergites. Small spinal tubercles on vertex
and abd. tergite VIII. Marginal tubercles small and low, irregularly present on some of
tergites II—IV. Dorsal hairs rather thick and stiff, with spear-shaped apices, spinally
on tergites I—III about 0.025 mm long; tergite VIII with 4 hairs 0.048—0.052 mm
long. Frontal tubercles smooth, well developed, with slightly diverging inner sides, at
their inner apices with 2 hairs of about 0.035 mm long, ventrally with 1—3 more hairs;
sinus frontalis conspicuously wide and with nearly flat bottom. Antennae about 11/15
times as long as body; basal segments brownish, slightly darker than the head; flagellum
evenly darker to blackish brown from near base to apex; segment III finely imbricated
from base to apex, near base with 2—3 hardly elevated, flattish rhinaria; processus ter-
minalis nearly 6 times as long as base of segment VI, with some 9—12 spiny hairs
besides the 4 on apex; segment III with about 22—28 hairs the longest of which is up
to 2/3 basal diameter of segment. Rostrum just reaching hind coxae as measured by
stylets; last segment with 14—16 hairs besides the 3 pairs near apex, 0.154—0.161 mm
long, nearly 115 times as long as second joint of hind tarsi. Legs about as pale as basal
half of ant. segment III, with the conspicuously bent apical 1/6—1/5 part of tibiae
blackish brown; femora on distal half ventrally, and near apex dorsally, slightly scabrous;
dorsal hairs at mid-length of tibiae 0.035—0.045 mm long, to as long as local diameter
of tibiae; first tarsal joints with 5, 5, 5 or 5, 5, 4 hairs, second joints ventrally hairy,
rather bluntly imbricated, and without spinules on imbrications. Siphunculi pale, apicad
gradually darker to brownish, distal 1/8 part rather dark brown; imbricated evenly and
superficially from base to near apex, with distal 1/9—1/8 reticulated; smallest diameter
on basal half about 0.050 mm, largest width in swollen part 0.061—0.065 mm, in the
reticulated area about 0.078 mm wide; flange thin, distinct. Cauda pale, elongated bluntly
triangular, not constricted, about 3/7 of length of siphunculi, with 7 hairs. Subgenital
plate with about 10—12 hairs along posterior margin, and 2 much longer ones on
anterior half.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Siph. Cau.
body III IV V VI

1 2.66 2.80 0.68 0.46 0.44 ORDEN 0:87 SUSE 3 0.71 0.32

2 2.54 ? 0.77 0.45 0.47 OISE 3072

(1—2, from Aquilegia formosa, Tioga Pass (3300 m.), Calif., U.S.A., 17.VIL’73, leg.
D. Voegtlin no. 75).

Discussion. Masonaphis (Ericobium) corylina (Davidson, 1914) was only known as
alatae, from Corylus sp., but Mason (1925) writes that according to Davidson the aphid
is common on Physocarpus capitatus. That is a most unusual host combination for a
Masonaphis. It is not clear whether the alatae developed on Corylus or had landed there.
Notwithstanding much searching no Masonaphis could later be found on Corylus or
Physocarpus.

At my request Mr. David Voegtlin searched Aquilegia formosa for more specimens
of Megourina lagacei spec. nov. He collected some larvae of that species, and some larvae
of Kakimia aquilegiae (Essig), but also two adult, slightly damaged apterae and two

126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

larvae of a Masonaphis (Ericobium). With the key in MacGillivray (1958) one arrives
at or near Masonaphis (Ericobium) corylina and my Voegtlin specimens may well
belong to that species. This can only be decided when alatae have been collected from
Aquilegia, or apterae viviparae from Corylus.

Masonaphis magna spec. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens body elongate oval, about 4.00—4.50
mm long. Tergum somewhat sclerotic but evenly faintly yellowish pigmented, not quite
smooth. Marginal tubercles usually present on tergites II—IV, inconspicuous, 0.007—
0.013 mm in diameter, rather flat; spinal tubercles of the same size common on the head
and tergite VIII. Dorsal hairs stiff, blunt, spinally on tergites I—III about 0.035 mm

ir DALE, UE

CER CHEN ite
oe A EE paid
ae Ae

Fig. 9. Masonaphis magna spec. nov. Apt. viv. fem.: a, antenna; d, last rostral segment; e, tip tibia
+ hind tarsus; f, siphunculus. Al. viv. fem.: b, antenna; c, cauda; g, siphunculus. a, b, c, d, f and g,
Pe UB NIOZ

D. Hire Ris LAMBERS: On American aphids 127

long, about 12 per segment; tergite VIII with 6—8 hairs up to 0.065 mm long. Frontal
tubercles with a few spinules on underside, diverging, with rather straight inner sides,
with 3—4 hairs; sinus frontalis bluntly V-shaped; median frontal process not developed;
antennae very long, to over 1 1/3 times length of body, pale brownish yellow, with the
very junctions between segments III/IV and IV/V dark brown, distal part of segment V
and all of VI dark brown; segment III evenly imbricated from base to tip, with in the
available specimens 5—8 rather evenly spread, hardly elevated, not bulging rhinaria in a
line on basal 1/3—2/5 part; processus terminalis 5—6 1/4 times as long as base of segment
VI, with no less than 12—20 spiny hairs besides the 4 on apex; segment III with some
30 blunt hairs, up to 0.022 mm long, about half basal diameter of the segment. Rostrum
short, reaching to middle coxae as measured by stylets; last segment very hairy, with
26—30 hairs besides the 3 subapical pairs, 0.190—0.205 mm long, to 12/3 times as
long as the small second tarsal joints of hind legs. Legs long, slender, pale, with the
conspicuously bent apices of the tibiae blackish brown; femora dorso-apically and laterally
somewhat imbricated; longest ventral femoral hair 0.045 mm long; dorsal hairs halfway
hind tibiae about 0.045 mm, about 2/3 of local diameter of tibiae; apices of all tibiae con-
spicuously hairy, with quite acute hairs; first tarsal joints with 5, 5, 5 hairs, second
joints hairy, sharply imbricated with rather distinct spinules on the imbrications. Siphun-
culi pale with conspicuously blackish apical 1/12—1/10 part, about 1/ of length of body,
imbricated from near base with apical 1/10 part distinctly reticulated, at base about 0.13
mm, from there gently tapering to halfway their length, to 0.075—0.080 mm, increasing
in diameter to a maximum of 0.088—0.092 mm, with the dark, reticulated apex about
cylindrical, much attenuated, only 0.050—0.055 mm wide; flange small. Cauda pale,
about half the siphunculi, with 8 hairs, one of which is dorso-apical; shape not describable
in available specimens. Subgenital plate with 16—20 hairs along posterior margin, two
longer ones near anterior margin and 1—4 hairs on the disc.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Siph. Cau.
body III IV V VI on III

1 4.22 4.75 145 0.92 0.85 O25 E32 5&6 1.09 0.48
2 4.19 5.78 1219 1.20 1.05 0.231.777, 8&8 1.19 0.64

(1, from Composite, 4 mi. E. of Sturgeon Heights, Alberta, Canada, 14.V1.’66, leg.
A.P.G. no. F 66—7—14 a; 2, from Composite, Mobley Lake, British Columbia, Canada,
12.V11.’66, leg. F.G.A.—A.P.G. no. F 66—7—12b).

Alate viviparous female.

Colour in life unknown. In mounted specimens body even larger than in apterae,
4.30—4.55 mm long; head and pronotum quite pale with some brown on the medial
side of the ocelli, mesonotum light brown, abdomen not pigmented. Antennae very long,
with segment I vaguely brownish, segment II and base of segment III basad rhinaria
pale, rest dark brown to brownish black, but segment IV on basal half lighter than
segment III; segment III with 17—19 rhinaria similar to those in apterae, very nearly
all in line, on basal 34 part. Legs as in apterae but femora dorso-apically brownish. Wings
with normal venation, veins brown, not bordered, but posterior margin of pterostigma

128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

with a brown border contrasting to the much paler stigma. Siphunculi much thinner than
in apterae, on basal half down to 0.050 mm, in the swollen portion up to 0.075 mm,
and in the reticulated 2/15 part down to 0.055 mm. Cauda with 7—9 hairs, flattened
in the slides. Other characters as ín apterae viviparae.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Siph. Cau.
body III IV V VI on III

1 4.34 5.56 1.26 1:18 1.00 0.30 + 1.54 18&19 1.05 0.51
2 4.53 5.58 1.14 1.03 1.00 037164 9172821189 4105 0.56

(1, with aptera no. 1; 2, from Composite, Lac La Hache, British Columbia, Canada,
6.V11.’66, leg. F.G.A. no. F 66—7—6 t).

Discussion. From the fact that this aphid was collected on one trip at three different
localities, it would seem to be fairly common. Unfortunately the Composite on which
all specimens were found was not identified.

It is very difficult to decide to which subgenus of Masonaphis H.R.L. this species
belongs. The imbrications on its second tarsal joints are about as spinulose as they are
in M. lambersi MacGillivray which one would classify as a Masonaphis sensu stricto.
No such aphid has been recorded from Compositae before. It is certainly not a member
of subgen. Oestlundia H.R.L. (pale head and thorax, no ornamentation in wings) or
Amphorophorina MacGillivray (arrangement of rhinaria). This leaves Masonaphis sensu
stricto and Ericobium MacGillivray. With MacGillivray’s (1958) key to Masonaphis
sensu stricto one could arrive at M. rhokalaza (Tissot & Pepper), but that species has
dark siphunculi, and fewer hairs on the last rostral segment. In her key to subgen.
Ericobinm one arrives at M. phacelia Essig, but that species has much less reticulation
to the siphunculi, long dorsal hairs, and a virtually smooth antennal flagellum. The tarsi
are very small. The hind tibiae are about 28 times as long as the second tarsal joints
(0.122 mm).

Because of its very large size, hairy last rostral segment, small tarsi and clear wings
this aphid should be easily recognizable.

Types. Holotype: alate viviparous female (measurements no. 1) from Composite,
4 mi. E. of Sturgeon Heights, Alberta, Canada, 14.VII.'66, leg. A.P.G. no. F. 66—7
— 14 d. Paratypes: aptera vivipara with data as for holotype; alate viviparous female,
from Composite, Lac La Hache, British Columbia, Canada, 6.V11.’66, leg. F.G.A. no.
F 66—7—6i; apterous viviparous female, from Composite, Mobley Lake, British
Columbia, Canada, 12.VII.’66, leg. F.G.A.—A.P.G. no. F 66—7-12 b.

Megourina gen. nov.

Type-species: Megourina lagacei spec. nov. Very similar to Megourella H.R.L., 1949,
but differing by: 5 or 4 hairs on the first tarsal joints instead of 3 hairs; median frontal
process distinct; sclerites in alatae obsolete on anterior abdominal tergites (always?);
yellow instead of black tibiae; marginal hairs on processus-like sclerites, like other dorsal
hairs with slightly capitate, very transparent tips.

D. Hire Ris LAMBERS: On American aphids 129

Discussion. The type and single species was first considered to belong to a subgenus
of Megourella, but there is no conceivable phylogenetic connection with that genus.
Therefore the strong resemblance of M. lagacei to the two Megourella species must be
considered convergence. The hairy rostrum, the structure of the dorsal hairs and the
shape of the front suggest relationship to the Nearctic Kakimia’s, associated with Saxi-
fragaceae, Scrophulariaceae and also Aguilegia.

Megourina lagacei spec. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens body oval, about 2.50—2.95 mm
long, about 1.40 mm wide. Tergum membranous, with dark brown scleroites about
0.016—0.050 mm in diameter at the bases of the spinal and pleural hairs, with very
strongly convex, processus-like, dark brown marginal sclerites, well developed ante-
siphuncular and rather smaller postsiphuncular sclerites, and a narrow, lighter brown
spinal bar across tergite VIII. Distinct semiglobular marginal tubercles, 0.006—0.013 mm
in diameter, singly, or sometimes in pairs or complex, regularly present on each marginal
sclerite on tergites II—V. No spinal tubercles present. Dorsal hairs on strong bases, thick,
stiff, dark, with seemingly finely furcated apex, but actually with a colourless thin knob,
spinally on abdominal tergite III about 0.035—0.055 mm long, the 4 hairs on tergite
VIII thinner and up to 0.065 mm long; marginal sclerites with 2—4 hairs each; about
14—16 hairs on each of abdominal tergites I—IV, of which the innermost pairs have
much smaller scleroites at their bases than the others. Head pale brown, smooth, with
only on the upper apex of the large, parallel, frontal tubercles a few nodules or spinules;
median frontal process seemingly absent, somewhat downwards. Antennae about as long
as body; first segment distinctly darker than head; segment III slightly imbricated,
blackish brown with pale basal portion and often also with a part distad the rhinaria
lighter, with 24—40 small, and sometimes some very small, slightly elevated, bulging
rhinaria irregularly arranged along 2/3—4/5 part of the segment, about twice as long
as segment IV; the latter on basal part mostly lighter than apical part of segment III,
distally brownish black like the more distal segments, without rhinaria; processus ter-
minalis 5—51/ times base of segment VI, with 7—10 small spiny hairs besides the 4
at the top; hairs on segment III about 14—18 in number, stiff, the longest one as long
as basal diameter of the segment. Rostrum (according to stylets) reaching beyond middle
coxae; last segment about 0.150—0.160 mm long, about 1 2/5 times as long as second
joint of hind tarsi, conspicuously hairy, with some 16—20 hairs besides the much longer
3 subapical pairs. Legs rather long; femora with blackish brown distal part, gradually
paler from there to base; tibiae pale to brownish yellow with blackish brown apices;
first tarsal joints mostly with 5 hairs, the slightly stouter middle one the shortest, but
apparently sometimes on one or two legs with 4 hairs, one of the lateral ones missing;
tarsal joints rather small, imbricated. Siphunculi blackish, about 1/5 of length of body,
on basal part with a few single spinules, remainder smooth, with a slender cylindrical
basal 1/3 part about 0.055 mm wide, distad gently increasing in diameter to a maximum
of about 0.070—0.075 mm, and evenly decreasing to a tip which is almost exactly as
wide as the thinnest part on basal half; flange small, about 11/7 times as wide as the
slightly incised part just below. Cauda dark, about 3/5 of the siphunculi, rather thick,
slightly tapering over 4/5 part, then rather suddenly tapering to the blunt apex, with

130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

probably 5—7 normal hairs (sometimes one hair of the lateral 3 pairs, sometimes the
dorsal-apical hair absent). Subgenital plate with 8—12 hairs along posterior margin, 2
longer hairs on anterior part, but also 8—10 hairs on the middle portion.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Siph. Cau.
body III IV V VI on III
i 2.89 2.96 0:92 0.44 0.43 OSEO SIN 3216 580057 0.31
2 2.69 2.61 0.82 0.38 0.38 0.15 + 0.770024 &274m052 0.29
3 2.68 2.66 0.79 0.39 0.36 Olst 073002618227 5092 0.27
4 2.63 2.83 0.86 0.42 0.39 0.15080" "26613007 0:53 0.30
5 2.80 3.03 0.95 0.43 0.44 0.1777540.820032:85 294.093 0.30
6 2.81 2075 0.85 0.40 037 0-15 0731002538027. HOS 0.29

(1—6, from Aguilegia formosa, Plaskett Creek, Highway no. 1, Monterey Co., Cali-
fornia, U.S.A., 5.IV.'61, leg. C. Lagace no. 66—IV—5 a).

Alate viviparous female.

Colour in life unknown. In mounted specimens head and thorax blackish brown;
abdomen almost without scleroites except at bases of spinal hairs of tergites V—VIII,
but with larger marginal sclerotic processes than in apterae. Antennae very dark with
only base of segment III pale; rhinaria on segment III except at the very base larger than
in apterae, and covering most of the circumference of the segment up to its top. Legs
much darker than in apterae. Wings with normal venation, the veins dark, not bordered.
Other characteristics much as in apterae, but siphunculi and cauda slightly more slender.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Siph. Cau.
body III IV V VI on III

1 2:95 2.99 0.92 0.44 0.44 0.15 + 0.82 49&55 0.53 0.27
2 2.64 2.80 0.85 0.43 0.39 0.15 + 0.77 47&47 0.46 0.24

(with apterae nos. 1—6).

Discussion. This aphid, according to specimens in the Essig collection, has been
collected several times before in California, from Agwilegia formosa, which host plant
it shares with Kakimia aquilegiae (Essig). Since it greatly differs from all other Nearctic
aphids, and its resemblance to the European Megourella spp. was discussed and p. 27,
no further discussion is necessary.

The species is dedicated to the late Mr. C. Lagace, who collected this and many other
fascinating aphids.

Types. Holotype: apterous viviparous female (measurements no. 1), from Aquilegia
formosa, Plaskett Creek, Highway no. 1 (Monterey Co.) California, U.S.A., 5.IV.'66,
leg. C. Lagace no. 66—IV—5 a. Paratypes: apterae viviparae and two alatae with
collecting data as for holotype. Some paratypes in the Lagace collection, Albany,
California, U.S.A.

D. HiLLE Ris LAMBERS : On American aphids 131

Fig. 10. Megourina lagacei spec. nov. Apt. viv. fem.: a, last rostral segment; c, antenna; d, posterior
part of abdomen; f, tip tibia + hind tarsus. Al. viv. fem: b, antenna; e, head; g, cauda; h, siphun-
culus. a and f, X 342; b, c, d, e, g and h, X 75

Nearctaphis argentinaeradicis (Gillette & Palmer, 1932)

According to the type material this apparently very rare aphid belongs in Nearctaphis
Shaposhnikov, and not in Aphis L. in which it was described.

The following items supplement the descriptions and figures by Gillette & Palmer
(1932) and Palmer (1952).

Apterous viviparous female. Head dorsally with about 24—30 hairs besides the 6—8
on the front, covered with blunt spinules or nodules. Last rostral segment to 0.18 mm,
with 4—6 hairs besides the 3 subapical pairs, 11, times as long as second joints of hind
tarsi. Dorsum of thorax and abdomen covered with little scleroites, bearing hairs, but
many hairs not on scleroites; tergite IV with 40—55 hairs, about 0.065 mm long, with

132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

extremely fine apices; tergite VIII with 12—16 hairs. Marginal tubercles present on abd.
segments IV, not on VI or VII, irregular in shape, and those on segment V larger
than all the others and composite, in dorsal view looking like an Eriosoma wax gland.
First tarsal joints with 4 hairs, 2 short and thick, 2 long and thin. Siphunculi with trans-
verse rows of bluntish spinules, with a rather wide flange.

With the kind permission of Dr. D. Fronk several paratypes collected 23.VIII.1930
and bearing the number 4830 were remounted and for the greater part returned to the
Palmer collection at Fort Collins, Colorado.

Neonasonovia inflata (Richards, 1962)

The species was described as a Hyperomyzus. But old apterae develop a vague brown
central sclerotic patch on the abdomen in combination with black siphunculi, while alatae
have an equally vague abdominal sclerotic pattern not at all like that in the four real
Hyperomyzus. Therefore I transfer Hyperomyzus inflatus Richards, 1962, to Neo-
nasonovia. Miss E. M. Patch recognized the species as new and gave a manuscript name
to specimens collected on 30.VII.’26 near Portland, Maine, U.S.A. on Hieracium, by
Mrs. I. M. Knight, and it was found in several localities in North Carolina by
C. F. Smith, J. O. Pepper and myself in 1963 and 1964. In the latter state it evidently
overwinters on Hieracium, for apterae, alatoid nymphs and alatae were found on
24.IV.'64 in two localities.

Neonasonovia nabali (Oestlund, 1886)

This species, described as Rbopalosiphum nabali from Nabalus albus, now Prenanthes
alba, is mostly treated as an Amphorophora Buckton in North American papers. Its
sexuals have not been recorded or described but I found them in very large numbers on
28.VIII.'56 in the Laurentide Park, Prov. Quebec, Canada, on the undersides of the
leaves of Ribes glandulosum. Alate males are brownish red, first instar oviparae greenish
yellow, after their first moult brilliant brownish red. Eggs at first very pale transparent
green, later black, on the branches.

In my hotel at Fredericton, N.Br., Canada, I transferred alatae from Prenanthes sp.
to leaves of Ribes nigrum and very soon obtained similar larval oviparae, and it would
seem that Ribes spp., at least R. glandulosum, are primary host plants of this species
which I transfer to Neonasonovia H.R.L.

Amphorophora fronki Knowlton, 1945, described from “Wild Currant” (= Ribes sp.)
is extremely nearly related to Neonasonovia nabali and congeneric. I could remount
some apterae from the type sample, one of which, though full of embryos, has a few
pseudosensoria on one atrophied hind tibia. I found up to 23 rhinaria on antennal
segment III, much fewer than in »abali, in which species the apterae from Prenanthes
alba have mostly about twice that number of rhinaria, which are smaller in size, in
addition to several on segment IV and sometimes V. But such a difference in number
of rhinaria is, in Neonasonovia and related genera, common between spring forms on
the primary host plant and summer forms from the secondary host plants. The swelling
of the siphunculi is similar, and rostrum, cauda, chaetotaxy, etc., do not show differences.
Only the large pigmented spinal scleroites of N. nabali from Prenanthes cannot be found
in remounted N. fronki, a case refound in N. picridis (Börner & Blunck), the type-
species.

D. HiLLE Ris LAMBERS: On American aphids 133

However, the number of rhinaria in alate N. fronki is certainly much lower than that
in gynoparae of N. nabali. Besides, an ovipara was found on Ribes in July. Therefore
nabali Oestl. and fronki Knowlton must be considered different taxa as long as no
authentic spring forms from the primary host, Ribes glandulosum, of Neonasonovia nabali
are known.

Probably Amphorophora petiolaris Knowlton & Allen, 1945, described as spring alatae
from Ribes petiolaris, Washington, also belongs to this group. I have not seen authentic
material. The host plant occurs in Utah. The name pettolaris may have priority over
fronki.

Neonasonovia nigra (J. McVicar Baker, 1934)

Julia McVicar Baker's aphids have received very little attention. Therefore it seems
useful to publish a more modern description of one of her species, originally described
as an Amphorophora from Stevia pubescens, from Mexico. She mentioned about 30
rhinaria on ant. segment III, 18 on IV, 4—5 on V, some more than in my material.

Apterous viviparous female.

Colour in life, according to Baker's description, very shiny black. In mounted
specimens body oval, about 1.30—1.55 mm long, about 0.60 mm wide, strongly convex.
Tergum with a solid, black, smooth shield extending from the suture-less fused tergites
of metathorax to abdominal segment V, and reaching down laterally to between the
stigmal pori; tergites VI—VIII with free sclerotic bars. No marginal or spinal tubercles
present. Dorsal hairs inconspicuous, blunt, only 0.004—0.008 mm long spinally on
abdominal tergites I—III; the 4—6 hairs on tergite VIII very little longer, to 0.010 mm,
about half as long as basal diameter of antennal segment III. Head frontally as dark as
abdominal dorsum, caudad paler; frontal tubercles not much developed, diverging, with
rounded inner sides, smooth, with 2 short hairs; median frontal process rounded, very
prominent, nearly half as high as the frontal tubercles. Antennae about as long as body
or slightly longer, blackish with only the very base of segment III pale; segment III near
base distinctly imbricated, more distally very lightly imbricated, along one side over its
whole length with 16—26 slightly elevated, a little bulging rhinaria, not in line but
nearly so; segment IV with 0—10 rhinaria; processus terminalis 31/,—5 times as long
as base of segment VI, with 2—4 small spines besides the 4 at apex; hairs on segment
III inconspicuous, to about 0.005 mm long, only 1/5—1/4 of basal diameter of the
segment. Rostrum reaching halfway between middle and hind coxae, last segment about
0.105—0.115 mm long, about 1 1/3 times as long as second joint of hind tarsi, with some
7—9 thin hairs of 0.013—0.021 mm besides the 3 pairs near apex. Legs in more pig-
mented specimens most conspicuous because basal 1/3 part of tibiae is pitch black, much
darker than any other part of body or appendages; femora blackish brown, paler brown
towards base; longest ventral femoral hair about 0.09 mm long; tibiae with middle 1/3
part pale to yellowish brown, at apex paler than on basal part; dorsal hairs halfway hind
tibia inconspicuous, to 0.009 mm long; first tarsal joints with 3, 3, 3 hairs; second
joints distinctly imbricated. Stigmal pori of abdominal segments I—V open, nearly
circular. Siphunculi blackish, smooth except for distinct striation in the annular incision
below the flange, 2/9—1/4 of length of body, strongly swollen; at base about 0.070 mm
wide, just above 0.050 mm, then widening to about 0.090 mm halfway their length and
gradually decreasing in width to 0.028 mm in the incision below the flange of about

134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Fig. 11. Neonasonovia nigra (J. M. Baker). Apt. viv. fem.: a, posterior part of abdomen; b, tip tibia
+ hind tarsus; c, last rostral segment; d, head; e, antenna. a, d and e, X 72; b, and c, X 342

0.037 mm. Cauda pale, slender, narrow-conical with a mostly distinct constriction, 6/11
—2/3 of the length of the siphunculi, with 5—6 normal hairs (mostly with a dorso-apical
one). Subgenital plate with 8—10 short, blunt hairs along posterior margin, with 2 hairs
of up to 0.020 mm near anterior margin.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. on Siph. Cau.
body III IV V IV III IV

1 152 1.48 040 0.20 0.19 0.09+0.46 19&20 O&1 0.53.0718

2 1:46 1.40 0.387 0.23, 0:20 0.10+0.35. 77168197 283 720537720.20

3 1.43 1.41 0,397 70:21 0.20 0.10+0.36 18&21 7&3 0.34 0.18

4 1.52 1555 0:45 0237 10:22 70.105040 722826 ESE ONO SONORO

(1—4, from Stevia glandulosa, 62 mi. E. of Villa Union, Sinaloa, Mexico, 1.11.'64, leg.
R. van den Bosch no. 64—2—1 f).

Discussion. Four adult apterae without larvae were collected by Dr. van den Bosch.
They are classified with some reluctance as Neonasonovia although they have short hairs.
However, the shape of the front, antennae including their sensoriation, round, not
covered stigmal pori, siphunculi with annular incision below the flange are so typical
that on these characters a decision is easy. Besides, the hairs in N. inflata (Richards) are
not much longer, and N. hieraci (Börner) has a position more or less intermediate
between inflata and the type species, picridis Börner & Blunck, as to dorsal hairs.

Because of its extensive black sclerotisation recognition of this aphid is very simple.

Neonasonovia zirnitsi Hille Ris Lambers, 1952

The author (1952) described this species from Ribes alpinum and Euphrasia sp. from
Sweden. At that time the life cycle had not yet been confirmed by transfer experiments.

D. Hire Ris LAMBERS: On American aphids 155

However, in 1967 I received living specimens on Ribes alpinum, found near Denekamp,
Netherlands, by the Phytopathological Service, and emigrants successfully colonized
potted Euphrasia nemorosa on which several generations developed. At my request Mr.
van der Schaaf then searched Euphrasia in the Denekamp area and he found several
specimens on this plant on 28.VII.'67. Therefore Ribes alpinum indeed is the primary
host plant of N. zirzitsi H.R.L. and Euphrasia spp. are secondary hosts.

Prevost (1959) described Hyperomyzus boerneri from Euphrasia rostkoviana and
Euphrasia sp. from Austria and Switzerland. Morphologically her type material cannot be
distinguished from zirnitsi developed on Exphrasia. Because the type material contained
several oviparae and two males (and also an alate oviparous female), it was supposed
that boerneri was a good species. However, further searching by the present author in
Switzerland and Austria revealed that this aphid, which is very common on Euphrasia
in the Alps, in September produces masses of alatae, settling on Ribes alpinum and
producing larvae, and some males. I now suppose that colonies remain on the secondary
host and that some oviparae are produced there when the temperature has become too
low for flight initiation, as in, e.g., Myzus cerasi (F.). Therefore Hyperomyzus boerneri
Prevost, 1959, is considered a synonym of Neonasonovia zirnitsi H.R.L., 1952.

Müller (1960) described Hyperomyzus sobrinus as host alternating between Ribes
alpinum and Euphrasia officinalis. Nothing distinguishes the paratypes he most kindly
gave me from Neonasonovia zirnitsi H.R.L., of which Hyperomyzus sobrinus Müller,
1966 is considered to be a synonym.

From Iceland Stroyan (1960) described Hyperomyzus boerneri subspec. thorsteinni
from Exphrasia frigida. This aphid very strongly resembles zirnitsi, but apterae collected
in August have the processus terminalis only about 4 times as long as the base of segment
VI, instead of 6 or more times as long as in true zirnitsi. Since Ribes alpinum is not
mentioned in the Iceland Flora, thorsteinni has apparently been split off the dioecious
zirnitsi, and has evolved far enough to be considered a species, as I do, transferring
it to Neonasonovia H.R.L.

Pentamyzus acaenae (Schouteden, 1904)

Through the kindness of Dr. H. Weidner, Hamburg, Germany, I could mount the
pickled material of Rhopalosiphum acaenae Schouteden, 1904. This appears to be a
Pentamyzus H.R.L., easily separable from the type-species, P. graminis H.R.L., by the
wide flange to the siphunculi (0.048—0.052 mm) and the thin part (0.030—0.035 mm)

Fig. 12. Pentamyzus acaenae (Schouteden). Apt. viv. fem.: a, antenna; b, cauda; c, siphunculus.

All DS 72

just below, where some transverse striae are present, and by antennal segment III being
twice as long as segment IV. Since the original slides seem to have been destroyed, a
lectotype, partly from Acaena splendens, Uschuaia, Tierra del Fuego, Argentine,

136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

14.X1.'92, leg. H. Michaelsen no. 147, was selected and returned to the Hamburg
Museum. Some of the lecto-paratypes, with collecting data as for lectotype, are in the
author’s collection.

Pentamyzus falklandicus spec. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens integumentum faintly yellowish. Body
elongated oval, 2.40—3.00 mm long, just over twice as long as its greatest width (rather
flattened specimens). Thoracic tergites free, tortoise shell-like sclerotic, or rather with
elevated, very low flattish warts, but along the median line apparently membranous;
abdominal tergites I—VIII similarly sclerotic and mutually free, but the membranous
median divide ending at anterior margin of tergite V or perhaps sometimes VI. Very
inconspicuous marginal tubercles, probably flat, about 0.005—0.008 in diameter, very
irregularly present on abdominal segments II—IV; spinal tubercles regularly present on
vertex, irregularly on abdominal tergites VII and VIII where they are more elevated and
larger than the marginal tubercles. Dorsal hairs scarce, on tergites I—IV spinally in a
very flat trapezoid, about 14—16 per segment, spinally on tergite III to 0.022 mm long,
stiff and blunt; tergite VIII with 8—12 mostly slightly longer hairs, to 0.030 mm long,
sometimes in a double row. Head broad; frontal tubercles just indicated, slightly
protruding beyond the convex front, wrinkled but not spinulose, nodulose or scaly, with
1—2 hairs each. Antennae of 5 segments, pale, with segment V brown, 2/5—3/7 of
length of body; segment I on inner side bulging; segment III without rhinaria, super-
ficially bluntly imbricated, more than twice as long as segment IV; processus terminalis
1 1/2—1 6/7 times as long as base of last segment, with mostly 2 spiny hairs besides
the 4 near apex of which one is always placed more than its own length below the tip;
antennal hairs scarce, only 5—7 on segment III and these half as long as basal diameter
of the segment. Eyes normal. Rostrum thick, reaching the middle coxae; last segment
acute, almost rostrate, about 0.14 mm long, about 8/9 of second joint of hind tarsi,
with 4—6 hairs of about 0.040 mm besides the 3 subapical pairs. Legs rather short and
thick, pale with perhaps the tarsi darker; longest ventral femoral hair about 0.030 mm,
stiff, bluntish, dorsal hairs on hind tibiae more acute, halfway tibiae about 0.30 mm long,
about 2/3 of local diameter of tibiae; first tarsal joints with 3, 3, 2 or 3, 3, 3 hairs;
second joints hardly imbricated; empodial hairs about 1/2—3/5 of length of claws.

Fig. 13. Pentamyzus falklandicus spec. nov. Apt. viv. fem.: a, antenna; b, siphunculi; c, cauda;
d, last rostral segment; e, hind tarsus; f, head. All X 72

D. Hire Ris LAMBERS : On American aphids 137

Siphunculi pigmented like the legs, nearly smooth with some wrinkles near base and some
striae near apex, about 1/7—1/6 of length of body, with basal 1/3 part about cylindrical
and there about 0.05 mm thick, suddenly strongly swelling to a maximum width of about
0.09 mm and from there tapering to the thinnest part, about 0.045 mm, just below the
rather thick flange of about 0.055 mm. Cauda thickly elongated triangular with convex
sides, less than half as long as siphunculi and similarly pigmented, with 6—10 normal,
acute hairs in irregular arrangement. The three rudimentary gonapophyses unusually
large, with spiny hairs of up to 0.030 mm. Subgenital plate with 14—18 bluntish hairs
along posterior margin and 2—4 much longer and more acute ones on anterior part.

Measurements in mm.

No. Length Ant. Ant. segments Siph. Cau.
body III IV NY
1 2.72 1219 0.44 0.17 0.192 0:25 0.39 0.25
2 3.00 127 0.47 0.18 0.16 + 0.25 0.43 0.25
3 2.60 1.24 0.45 0.19 OMS 70: 27; 0.37 0.22
4 275 1.26 0.44 0.19 05028 0/39 0.23
5 2.81 1.57 0.49 0.21 0.1877.20:29 0.46 0.26
6 2295 1.26 0.44 0-19 0.195:720.28 0.39 0.23
i 2.47 1.18 0.42 0.18 0.14 + 0.26 0.37 0.23

(1—7, from Poa flabellata, Falkland Isles, 21.V1.’37, leg. A. G. Bennett no. 8246).

Discussion. The closest relative of this aphid is P. acaenae (Schouteden) supposedly
living on Acaena splendens, but there was also a piece of a blade of grass in tube no. 147.
In P. acaenae the processus terminalis is shorter, 1—1 1/3 times as long as base of
segment V; the siphunculi swell gradually from 0.048 near base to a maximum of 0.065
mm, from where they gradually narrow to 0.035 mm just below the thin flange. The
body in acaenae is very different in shape, e.g., 1.94 mm long with a greatest width of
1.21 mm.

The sample consisted of 13 apterae and one larva. The latter has smooth hind tibiae.
In one aptera the embryos can easily be examined. They have 5 antennal segments,
segment III being hairless. The last rostral segment has 4 hairs besides the 3 subapical
pairs, and it is distinctly rostrate. The chaetotaxy of the dorsum is not visible.

Types. Holotype: apterous viviparous female (measurements no. 2), from Poa
flabellata, Falkland Islands, 21.1V.’37, leg. A. G. Bennett. Paratypes: apterae viviparae
with collecting data as for holotype. Holotype and most paratypes returned to the British
Museum (Natural History); some paratypes in the author's collection.

Thripsaphis Gillette, 1917

Börner (1930, 1952), Quednau (1954), Eastop (1966), and Richards (1971) use a
subdivision of what Palmer (1952) has under Thripsaphis Gillette. This, in its most
recent form by Richards, is as follows:

a. Thripsaphis Gillette, 1917, type species Brachycolus ballit Gillette, 1908. Empodial
hairs spatulate.

138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

b. Allaphis Mordvilko, 1921, type species Callaphis caricicola Mordvilko, 1914 1).
Empodial hairs setiform, apterae viviparae without wax glands (“cribriform discs” of
Richards, 1971) or with wax glands not visible (Quednau, 1954).

c. Trichocallis Börner, 1930, type species Allaphis caricis Mordvilko, 1921. Empodial
hairs setiform, apterae viviparae with dorsal wax glands.

Eastop (1966) uses Allaphis and Trichocallis as subgenera of Thripsaphis. The sub-
division looks clear enough, but there are some objections. The case of Thripsaphis
utabensis Knowlton & Hall, discussed on p. 149, suggests that the presence or absence
of wax glands is not a very good character. But besides, a very close examination of those
species the four authors thought to have no wax glands, revealed that these are present,
though often only caudo-laterally on abd. tergite VIII, and in dorsal view hardly visible.
A more serious objection arises from the accumulation of misidentifications of the type
species of Allaphis and Trichocallis.

Through the kind help of Dr. G. Shaposhnikov, Leningrad, I could remount and
examine original material of “Callaphis” caricis and “C.” caricola Mordv., collected
from Carex in 1907 and labelled by Mordvilko. The results are:

a. Callaphis caricicola Mordvilko, 1914, later renamed Allaphis caricicola (Mord-
vilko, 1914) by Mordvilko (1921). Börner (1952) synonymized Thripsaphis thripsoides
Hille Ris Lambers, 1939 with caricicola Mordv. However, this is quite wrong. Actually
Thripsaphis gelrica Hille Ris Lambers, 1956 is a synonym of Callaphis caricicola Mord-
vilko, 1914, and that is a species with markedly spatulate empodial hairs. Therefore
Callaphis Mordvilko, 1909, type species Callaphis caricicola Mordvilko, 1914, and
Allaphis Mordvilko. 1921, nomen novum pro Callaphis Mordvilko, 1909 nec Walker,
1870, become synonyms of Thripsaphis Gillette.

b. Allaphis caricis Mordvilko, 1921, the type species of Trichocallis Börner. Thrips-
aphis thripsoides Hille Ris Lambers, 1939 is a synonym of Allaphis caricis Mordvilko,
1921, and not of Callaphis caricicola Mordv., as Börner and later authors thought.
Allaphis caricis Mordv. is a species with setiform empodial hairs, and, as Thripsaphis
thripsoides H.R.L. according to Richards (1971), without dorsal wax glands in apterae
viviparae.

As the presence or absence of wax glands is hardly a sensible character, I recognize
only two subgenera, Thripsaphis Gilette, 1917, and Trichocallis Börner. The fact that
Börner (1930, 1952) misidentified Aphis cyperi Walker, 1848 with Allaphis caricis
Mordv., and (1930, p. 165) erroneously synonymized his genus Trichocallis erected on
p. 127 of the same paper with Phyllaphoides Takahashi, fortunately has no consequences.
Also cyperi Walker has setiform empodial hairs, even if it has very many wax pores on
the dorsum.

1) The years of publication of the genus Callaphis Mordv. and the species caricis Mordv. and
caricicola Mordv. were dubious. Mordvilko (1921, p. 58) writes “caricicola Mordv. (1908)”.
Börner (1952, p. 282) could not find a reference to the species in Mordvilko’s papers of 1908.
Neither could I. Börner then writes that Mordvilko (1914, p. 27) first turns up with Callaphis,
with as single species C. caricicola Mordv., in a key to genera. But as Mordvilko treats both genus
and species as if they were described earlier by him, I continued the search, and found in a footnote
in Mordvilko’s (1909) paper on aphid biology on p. 102: “In die Gruppe Callipterina bewohnt
nur die Gattung Sipha Pass. Kräuter und namentlich Gramineen, sowie die Gattung Callaphis
Mordw. (caricicola Mordw. und caricis Mordw.) die Blätter von Carex;”. Consequently Callaphis
Mordv. should have 1909 as year of publication; caricicola Mordv. remains a nomen nudum till
1914; and caricis Mordv. till 1921. when it was described as an Allaphis.

D. Hire Ris LAMBERS : On American aphids 159

In Richards’ revision (1971) of the sedge aphids of the world a few species and one
genus have not been discussed. He lists Thripsaphis striata Bozhko, 1961 as a synonym
of Subsaltusaphis ornata (Theob.) which is not suggested by the photograph accom-
panying the description. Dr. Maria Bozhko most kindly made a type available for
examination. It appears that Thripsaphis striata Bozhko, 1961 belongs in Neobacillaphis
Huculak, 1968, type species Neobacillaphis szelegiewiczi Huculak, 1968, a species and
genus not mentioned by Richards. N. striata (Bozhko) differs from N. szelegiewiczi
Huculak by a shorter and broader body with paired, dark, spino-pleural sclerites; an
evenly rounded abd. tergite VIII (as in T. ossiannilssoni H.R.L.) which has a number
of small hairs on its disc as in N. szelegiewiczi; by shorter antennae without secondary
rhinaria in apterae, with a processus terminalis only half as long as the base of ant.
segment VI, with conspicuous, dense transverse rows of spinules from segment III—VI,
and with long hairs, mostly longer than the largest diameter of ant. segment III; by
having no umbrella-shaped, but inverted-wedge shaped dorsal hairs.

Also Thripsaphis caricicola Swain, 1918 is not mentioned by Richards (1971). The
type material of this California sedge aphid can not be found. At any rate the name is
not available, because it is a junior secondary homonym of Callaphis caricicola Mord-
vilko, 1914.

A number of new members of the subgenus Trichocallis are described hereafter. The
North American species key as follows:

KEY To THE NORTH AMERICAN SPECIES OF THE SUBGENUS Trichocallis

Apterous viviparous females.

1 (2) Median process on front as long as ant. segments I and II together. Carex
trichocarpa. Canada.

T. daviaulti Quednau

2 (1) Median frontal process shorter than ant. segments I and II together.

3 (4) Abd. tergite VIII with its caudal margin triangular with slightly convex sides
or with a faint process bearing two hairs. Resembles strongly caricis Mordv.,
but differs in pattern of dorsal nodules. Colorado, Quebec.

T. producta Gill.

4 (3) Abd. tergite VIII with caudal margin in the middle rounded or straight or
indented.

5 (16) Hind margin of tergite VIII, posterior corners of tergite VII, and often
various other places, e.g. around spinal and pleural hairs, and distal parts
of antennal segments, with stippled wax pores.

6 (11) Distance between the two middle hairs on posterior margin of abd. tergite
VIII only about 9/10—11/ times as large as the distance from either of
those hairs to the nearest marginal hair more basad. Tergite VIII always very
evenly rounded, more or less semicircular, often with a number of tiny hairs
on its disc. Group of T. osstannilssoni H.R.L.

7 (10) Ant. segment III only about 31/,—61/, times as long as its greatest width.
Dorsal ornamentation of abdomen consisting of long, wavy, here and there
coalescing, rows of very small semi-acute spinules.

140

8 (9)

9 (8)

10 (7)

11 (6)

12 (13)

13 (12)

14 (15)

15 (14)

16 (5)
17 (18)

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Disc of abd. tergite VIII with 6—15 mostly very inconspicuous, thin hairs
besides the single or partly double row of much larger marginal hairs. Only
Europe to Japan.
T. ossiannilssoni H.R.L. sensu stricto
Disc of abd. tergite VIII without hairs. California.
T. ossiannilssoni subsp. pacifica subsp. nov.
Ant. segment III at least 8, mostly 10 or more times as long as its greatest
width. Dorsal ornamentation consisting of short rows of 2—4 blunt nodules
mixed with half-fused single large nodules up to 0.009 mm in diameter.
Probably 7. ossiannilssoni X T. subverrucosa, found with the potential parents.
California.
T. hybrida spec. nov.
Distance between the two middle hairs on posterior margin of abd. tergite
VIII more than 11/ time as large as the distance from either of those hairs
to the nearest marginal hair more basad. Tergite VIII mostly not evenly
rounded, either with a blunt angle between sides and rounded apex (or, not
as a result of mounting, posterior marginal hairs far from posterior margin
of tergite), or with a straight or slightly concave part between the middle
hairs on posterior margin.
Distance between middle posterior hairs on margin of abd. tergite VIII
21/—31/ times as large as distance from either of these hairs to the nearest
marginal hair more basad. Only posterior angles of tergite VIII with stippled
wax glands, none in the middle of posterior margin. Antennae long, up to
3 times as long as width of head across and including the compound eyes.
Front with very pronounced median process of 0.065 mm long. California.
T. californica spec. nov.
Distance between middle posterior hairs on margin of abd. tergite VIII less
than 2 times distance from either of these hairs to nearest marginal hair more
basad. A broad zone of stippled wax glands along the margin of tergite
VIII, also in the middle. Median frontal process not exceeding 0.055 mm.
Row of hairs along posterior margin of abd. tergite VIII placed up to their
length inwards from posterior margin of that tergite. Dorsal ornamentation
virtually absent. More than posterior half of tergite VIII covered with stippled
wax glands. Large fields of stippled wax glands around the marginal, pleural
and spinal hairs of thorax and abd. tergites I—VII. Many States in North
America; Europe.
T. cyperi (Wlk.) sensu latiore
Row of hairs along posterior margin of abd. tergite VIII arranged on, or very
nearly on, that margin. Dorsal ornamentation very conspicuous, consisting
of short rows of 2—4 subacute spinules suggesting imbrication. Along the
the median line stippled wax glands on abd. tergite VIII only on posterior
1/4—1/3 of the segment. Fields around spinal hairs on abd. tergites IVI
consisting of 0—10 glands. Utah.
T. utahensis Knowlton & Hall
No wax glands visible or only a few on abd. tergite VIII.
The 10—16 hairs on abd. tergite VIII thin, only about 0.015—0.030 mm

D. Hire Ris LAMBERS: On American aphids 141

long. Dorsal ornamentation on abd. tergites II—V consisting of rather sharp,
small spinules, often in short rows of 2—5, but distance between rows, or
between isolated spinules at least 3 times diameter of spinule. Tergite at
posterior margin not indented in the middle. Idaho, California.
T. scabra spec nov.
18 (17) The 6—10, mostly 8, hairs on abd. tergite VIII stout and stiff, the longest
of the set 0.050—0.085 mm long, when rather short often somewhat blunted.
Dorsal ornamentation on abd. tergites I—III mostly consisting of more or less
flat, partly strongly transversely oval warts which may measure 0.010 mm in
one direction, and which are not farther apart than their greatest length.
Tergite VIII on posterior margin somewhat depressed or indented in the
middle. Many North American States, also Europe.
T. verrucosa Gill. sensu latiore

Thripsaphis (Trichocallis) californica spec. nov.

Apterous viviparous female.

In life very pale yellowish white with glassy, transparent margins, more or less shiny;
antennae dark; legs glassy. In mounted specimens body about 2.20—2.70 mm long,
maximum width about 0.37 of length, pale with in the darkest specimens a faint
browning along the very margin of abdomen, especially on abd. tergites VII and VIII,
and with more distinct very thin, brownish intersegmental transverse lines. Spinal hairs
on abd. tergite III 0.008—0.010 mm long; marginal hairs to 0.016 mm long; the 8—10
marginal, medium-thin, hairs on tergite VIII to 0.040 mm long, with middle posterior
pair about 2/,— 314 times as far apart as distance to nearest more lateral hair. Stippled
poriform wax glands clearly visible in the darkest specimens on their marginal pig-
mented areas, from tergites III to VIII; on the more anterior of these tergites around
the marginal hairs; in quite pale specimens at most visible on tergite VIII, but rather
frequently not visible, also because they may be on the edge of the body; legs and
antennae apparently without wax glands. Integumentum on mid-dorsum with irregularly
arranged, rather pointed nodules, shorter than their basal width of about 0.002 mm, in
transverse direction about 3—4 times their own width apart, but in longitudinal direction
much further apart; marginally these nodules are nearly absent, but where present they
appear to be inclined caudad; from tergite VI caudad the nodules change to very short
spinules in transverse rows of 2—3 on tergite VI, 2—7 on tergite VIII which looks
spinulosely imbricated. Head with a median band of small blunt nodules which, more
laterally, are much flatter, blunt, more disperse, and hardly visible. Front strongly, down-
wards, protruding with a rounded median process as long as ant. segment II with 2 pairs
of hairs of about 0.035 mm long. Antennae 3/5—2/3 of length of body, with segments
I, II, and about basal 1/3—1/2 of III pale, rest dark brown; flagellum with spinules on
the straight, fine, imbrications; segment III without rhinaria, much longer than segment
VI, with some 12—16 straight, acute hairs up to 3/5 as long as diameter of segment at
its constricted base, about 14—17 times as long as its largest width. Rostrum reaching
to half-way middle coxae; last segment half as long as second joint of hind tarsi, with
2 hairs besides the 3 subapical pairs. Legs as pale as body with only the second tarsal
joints infuscated; tibiae to near base inconspicuously spinulose except ventrally, hind
tibiae between 2/7 and 1/3 of length of body; apical tibial hairs not differentiated; first
tarsal joints very spinulose, with 5 ventral hairs; second tarsal joints 1/5 of length of

142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

hind tibia; empodial hairs fine. Porus of siphunculi about 0.035 mm in diameter, hardly
raised. Tergite VIII roughly in the shape of a rectangle with sides of 0.065 and 0.035
mm of which the upper corners are replaced by quarter circles with a radius of half the
short side; sides slightly convex, posterior margin faintly convex; disc without hairs.
Cauda knobbed, the knob distinctly longer than wide, with 13—16 short hairs.

Measurements in mm.

No. Length Ant. Ant. segments Cau.
body III IV V VI
1 2.66 172 052 0.34 0.33 0.182570.19 0.15
2 2.34 1.60 0.48 0.31 Ot 0.18 + 0.16 0.12
3 2.63 1.74 0:55 0.36 032 Onl Se ONS 0.13
4 2.67 177 0.53 0.35 0.33 OLLIEERONLS 0.13
5 295 1.57, 0.47 0.31 0.30 Ol Se OG 0.12
6 2.63 11.0) 0.48 0.34 0.32 OMS EEN 0.14

(1—6, from Carex sp., Inverness, Point Reyes, California, U.S.A., 22.11.’64, leg.
D.H.R.L. no. 148).

Discussion. The species might easily be mistaken for Thripsaphis verrucosa Gill.
without rhinaria on ant. segment III, but the structure of abd. tergite VIII is quite
different. In only a few specimens, the more pigmented ones, the wax glands are quite
distinct.

Because the presence or absence of wax glands is such an unreliable character, I have
made an effort to find other discriminants, such as chaetotaxy and the fine sculpture of
the integumentum, for species of this genus. Quite good slides are needed for studying
the sculpture of the integumentum.

The Carex on which this species occurred was a tall species with leaves of over 10 mm
wide, growing in a ditch. It was thought to be C. comosa, but inflorescences were not
available.

Types. Holotype: apterous viviparous female (measurements no. 6) from Carex?
comosa, Inverness, Point Reyes peninsula, California, U.S.A., 22.11.’64, leg. D.H.R.L.
no. 148. Paratypes: apterae viviparae with collecting data as for holotype.

Thripsaphis (Trichocallis) cyperi (Walker, 1848)

The most characteristic features of this species are the following: the marginal hairs
on abd. tergite VIII stand far inwards from the posterior margin of that tergite, there
are no hairs on the disc of this tergite, the antennae in adult apterae have secondary
rhinaria, and the empodial hairs are not spatulate but setaceous. T. caricicola (Mordv.)
is very similar, has the same exceptional arrangement of marginal hairs on abd. tergite
VIII, but has, in all specimens studied, some extra hairs on the disc of that tergite, has
no secondary rhinaria on the antennae in apterae, and spatulate empodial hairs. T. cyperi
is widely distributed in the Nearctic and Palaearctic, but T. caricicola is only known
from the Palaearctic Region.

T. vibei H.R.L., 1952, which has the hairs on abd. tergite VIII arranged as in

D. Hire Ris LAMBERS: On American aphids 143

T. cyperi, was described from Greenland, where also a form with longer antennae occurs.
Richards (1971) has listed T. vibei as a synonym of T. cyperi, but does not mention for
which reason. Because T. vibei does not show the very elongate shape of T. cyperi, and
has rather shorter antennae, it can easily be distinguished. In T. vibei the flagellum of
the antennae in apterae is 0.9—1.0 times the greatest width of the body. In T. cyperi
the flagellum is 1.4—1.8 times as long as the greatest width of the much more elongate
body, depending on the degree of flattening of the body through mounting. T. cyperi
subsp. arctica H.R.L., 1955, agrees with T. cyperi as to shape of body, but the absolutely
shorter antennae are more like those of T. vibei, and therefore I (1960) treated it as a
subspec. of T. vibei. In this subspecies the flagellum of the antennae in apterae is
1.15—1.25 times as long as the greatest width of the body.

As yet T. vibei is only known from two good samples from Greenland, T. vibei subspec.
arctica H.R.L. from one sample from Greenland, and seven samples from Iceland; it
produces sexuals in Greenland at the end of July, in Iceland in the beginning of August.
T. cyperi is known from the more northern States of U.S.A., e.g., Minnesota, and in a
broad belt across Canada at least as far north as Labrador, in Europe from west to east
reaching to the north as far as Northern Sweden, and to the south into the Alps; as far
as known its sexuals are produced in the autumn.

Thripsaphis (Trichocallis) hybrida spec. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens body 2.20—2.45 mm long, about
3 times as long as its largest width; seemingly evenly pale, with abd. tergite VII posterio-
laterally, and tergite VIII marginally and caudally smoky brown, but actually colourless
intersegmental lines between segments posteriorly bordered by pale brown visible on
abdomen. Tergum densely covered with flat-topped, small, oval warts of up to 0.006
mm in the longest, transverse axis, and these warts very often divided in two equal parts;
laterally warts much smaller and even pointed, and on tergites VII and VIII also medially
passing into spinulose imbrications; the ornamentation much as in T. verrucosa Gill., but
warts smaller. Dorsal hairs on tergite III about as long as the width of the nearest wart,
cephalad of the siphunculi marginally not longer, but tergite VII (and mostly VI, some-
times V) marginally with stout, sharp hairs of 0.030—0.055 mm; tergite VIII, only
marginally, with 8—11 slightly longer, stout hairs of which the middle posterior pair
stands not or hardly farther apart than the other hairs; no hairs on the disc of tergite
VIII. Wax glands indistinctly present on posterior margin of tergite VIII. Head with
small, blunt nodules; front strongly protruding but not quite as far as the length of
ant. segment II, with 2 stoutish sharp or blunt hairs 0.040—0.045 mm long; more
laterally and caudad two shorter hairs on low processes. Antennae with segments I, II
and basal part of III pale, rest dark brownish; flagellum imbricated with very short
spinules on the imbrications; segment III about 9—12 times as long as its largest width,
without rhinaria and glands, just longer than segment VI, with 6—9 hairs up to half
as long as diameter of segment at its constricted base. Rostrum not nearly reaching middle
coxae; apical segment about half as long as second joint of hind tarsi, with 2 short hairs
besides the 3 subapical pairs. Legs pale, with tarsi distad brownish; tibiae especially
dorsally spinulose from base to apex; hind tibiae 14 of length of body; apical tibial hairs
not differentiated; first tarsal joints with 5 ventral hairs; second joints less than 14 of

144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

length of hind tibiae; empodial hairs setaceous. Siphunculi slightly elevated, on the
margin of tergite VI which is densely covered with small, just blunt, nodules. Tergite
VIII semicircular to semioval, evenly rounded also on posterior margin, about 4/7 times
as long as its basal width, with anterior middle portion pale, rest dark smoky brown.
Cauda pale, the knob trapezoid, as long as wide, with 14—16 short hairs.

Measurements in mm.

No. Length Ant. Ant. segments Cau.
body III IV V VI

1 2.44 1.14 0.32 09 0.19 0.13 + 0.14 042
2 2.21 0.97 027 0.16 0.15 04227012 0.11

(1—2, from Carex sp., Wheeler Springs (Ventura Co.), California, U.S.A., 19.V.'61,
leg. R. van den Bosch, no. 61—V—19 r).

Discussion. The two specimens of T. hybrida were part of a rather large sample, no.
61—V—19 r, collected by Dr. R. van den Bosch. In the sample the following different
apterous forms could be distinguished.

1. T. ossiannilssoni pacifica, described on p. 145.

2. T. verrucosa Gill., one quite typical specimen with rather large, flat warts from
pronotum to abd. tergite VI, a distinct flat furrow between two scabrous protuberances
on the posterior margin of abd. tergite VIII, and some rhinaria on segment III of the
long antennae.

3. T. hybrida, two specimens with small warts mostly in pairs suggesting small trans-
versely oval warts, an evenly rounded tergite VIII with traces of stippled wax glands on
N.E. and N.W. part near the margin, and short antennae without secondary rhinaria.

4. Specimens with the dorsal ornamentation exactly as in T. hybrida, but with
abdominal tergite VIII rather as in T. verrucosa, with the flat furrow in the middle of
posterior margin, and the two stout hairs beside it on a scabrous, very little pronounced,
elevation; antennae without secondary rhinaria.

5. Specimens much like no. 4, but covered with very small nodules in transverse
rows, seemingly without a trace of warts, except on the middle of mesonotum where they
appear to be subdivided. Antennae without secondary rhinaria.

I should add that in nos. 2—5 the hairs on tergite VIII are all stout as in typical
T. verrucosa.

One wonders what happened on this plant. Are these various types the products of
hybridization? With that in mind I proposed the name T. hybrida for no. 3. They are
like an intermediate hybrid between T. ossiannilssoni, and nos. 4 and 5, which I describe
on p. 151 as T. verrucosa subverrucosa subspec. nov.

It should be emphasized that the differences in the structure of abd. tergite VIII are
not artifacts. All the specimens were remounted and regained their shape completely in
water, and kept it subsequently in rather thick slides.

Types. Holotype: apterous viviparous female (measurements no. 1), from Carex sp.,
Wheeler Springs (Ventura Co.), California, U.S.A., 19.V.’61, leg. R. van den Bosch
no. 61—V—19 r. Paratype: aptera vivipara with collecting data as for holotype.

D. Hire Ris LAMBERS : On American aphids 145
Thripsaphis (Trichocallis) ossiannilssoni pacifica subsp. nov.

Apterous viviparous female (from one specimen).

Colour in life not known. In mounted specimen body about 3 times as long as its
greatest width; head pale; thorax laterally with smoky, sclerotic areas; abdomen with
caudad larger and darker marginal sclerites on each segment, and with very vague, paired,
widely separated, short, spino-pleural sclerotic bars which on tergite VII are fused to one
bar, and very nearly so on tergites V and VI; sclerotisation slightly more distinctly
pigmented on posterior tergites; tergite VIII about as dark as marginal sclerites on tergite
VII. Hairs on dorsum very short, spinally on tergite III about 0.004 mm long, marginally
about 0.009 mm; tergite VIII only along its margin with 12 much stouter, acute hairs
0.030—0.065 mm long, but no hairs on its disc. Wax glands of the stippled pore variety
only visible around the marginal hairs of abd. tergites I—VI, over most of the marginal
sclerites of tergite VII and on most of tergite VIII except the middle anterior part; I
cannot find them on antennae or legs. Dorsum only distinctly spinulose where it is
pigmented, the spinules very small, not longer than wide at base, and rather blunt, in
rather short rows, rather far apart. Head seemingly smooth because it is pale; front
protruding about as far as ant. segment II is long. Antennae darker than head; segment
III about 61/4 times as long as its maximum width, without rhinaria; segment VI consid-
erably longer than III; flagellum faintly imbricated with rather inconspicuous and widely
spaced spinules on the imbrications; longest of the very few hairs on segment III half
as long as diameter of segment at its much constricted base. Rostrum reaching the middle
coxae; last segment with 2 hairs besides the 3 subapical pairs, about 4/7 of second joint
of hind tarsi. Legs rather short, pale with bases of tibiae slightly darker than apices;
tibiae on distal half slightly spinulose, very faintly so dorsally on basal half; hind tibiae
about 14 length of body; apical tibial hairs not differentiated; first tarsal joints with
5 ventral hairs; second joints about 1/, the length of the corresponding tibiae; empodial
hairs fine, setaceous. Siphunculi nearly poriform, not measurably elevated, on the
marginal sclerite of abd. tergite VI. Tergite VIII semicircular, caudally hardly flattened.
Cauda with strongly transverse knob, with 13 rather short hairs.

Measurements in mm.

No. Length Ant. Ant. segments Cau.
body III IV V VI

1 1.74 05260: 50.17 0.12 0.13 Os 20E 0.08

(1, from Carex, Wheeler Springs (Ventura Co.), California, U.S.A., 19.V.’61, leg.
R. van den Bosch 61—V—19 r).

Discussion. Thripsaphis ossiannilssoni H.R.L. is now known from the Netherlands to
Japan (Higuchi, 1972), and specimens from various European countries hardly differ
from Japanese material. All have a number of often very inconspicuous hairs on the
disc of abd. tergite VIII besides the much longer and thicker hairs along the margin.
Mostly one sees only the bases of these hairs as perforations in the dark tergite. However,
in the Californian specimen there is no trace of such hairs.

T. ossiannilssoni can easily be recognized with the key on p. 139-141.

146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Type. Holotype: apterous viviparous female, from Carex sp., Wheeler Springs (Ventura
Co.), California, U.S.A., 19.V.’61, leg. R. van den Bosch no. 61—V—19 r.

Thripsaphis (Trichocallis) producta Gillette, 1917

Under this name Gillette (1917) and Palmer (1952) very clearly described, or figured
part of, the species which Richards (1971) described as Allaphis thripsoides Hille Ris
Lambers. Under Allaphis thripsoides, Richards writes that Thripsaphis foxtonensis
Cottier, 1953, is a synonym and he quotes also Eastop (1966) in this context. Eastop,
however, separated the two species. They are closely related but totally different. The
types Dr. Cottier gave me show in the apterae antennae of 1.80 mm long, fully twice
as long as in average thripsoides = caricis Mordv., and nearly reaching to abd. tergite VI.
Also the alatae are very different, by having broad, black sclerotic, quite free bars across
the abdomen instead of a solid sclerite like T. caricis Mordv. This is clearly shown in
Cottier’s figures.

The confusion about 7. producta appears to be that the type-specimen in the U.S.
National Museum does not agree with the original and Palmer’s (1952) description. !)
This is not very surprising. Not until 1932 did Gillette and Palmer mention types for
their species, and the unfortunate mix-up with T. balli is clearly shown in Gillette &
Palmer (1931) where the VIIIth abdominal tergite of ball; is drawn over the caption
T. producta.

American T. producta is not the same as European T. caricis, though the two are
extremely alike. There is a consistent difference in the pattern of the nodules on the
abdominal tergum. In T. caricis they are small on the middle of the body, often subacute,
and invariably arranged in somewhat wavy, often coalescing, very long transverse rows.
In T. producta from America the nodules are larger and if there are rows, they consist
of 2—3 nodules.

T. thripsoides = caricis, and T. producta are said by Eastop (1966), and consequently
Richards (1971), to have no wax glands, or cribriform discs. They are clearly visible in
specimens, especially near-adult larvae and apterae viviparae, with a dark eighth abd.
tergite, along its sides, and in larvae also on the marginal sclerites. In pale adults they
are invisible.

1) The type slide, borrowed from the U.S. National Museum, Washington, D.C., (Cat. no. 41948)
contains an alate and an aptera vivipara both indicated as type, and an alate indicated as paratype.
All are indeed Thripsaphis ballii (Gillette) as stated by Richards (1971) and confirmed by Quednau.
It has very clearly been relabelled in the handwriting of ? C. P. Gillette. Under the right - hand label
is another label on which the original name, which I could not read, ís obliterated by a narrow strip
of paper on which the name “T. producta n. sp.” is written. The specimens were found “on Carex
sp. / 6-3-1915 / Ft. Collins, Colo. / Coll. L. C. Bragg / Colo. Agr. Exp. Sta. [Ac. 1005”. Another
slide labelled metatype, with one label “Thripsaphis / producta n.sp. / on Carex / 8.17.07 / Ft.
Collins, Colo / C. P. G. Coll”, contains 6 apterae viviparae and two larvae which are in excellent
condition and agree with the original description. It is quite clear that an error in labelling has been
made when about 1932 types were indicated by C. P. Gillette and/or M. A. Palmer.

I made a formal proposal to the International Commission on Zoological Nomenclature to approve
the designation of a lectotype of Thripsaphis producta Gill. that agrees with the original description.

It is the lowest specimen on the right, on slide: U.S. Nat. Museum no. 73040, labelled on the
left: “Thripsaphis / producta / Gilette, 1917 / lectotype (at arrow) / and paratypes / D. Hille Ris
Lambers, 1973”. On the right labelled as mentioned above for the metatype.

D. HiLLE Ris LAMBERS: On American aphids 147

Thripsaphis (Trichocallis) scabra spec. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens body evenly yellowish, 2.40—2.90
mm long, about 21, times as long as its maximum width which is on the posterior third
of the body. Spinal hairs on abd. segment III thick, inconspicuous, about 0.007 mm long,
marginal hairs 0.009 mm, all sharply curved backwards; the 11—16 hairs on tergite VIII
mostly strongly curved near the middle, the longest (median posterior pair) 0.022—
0.034 mm long, the shortest 0.013 mm long; often a few hairs on tergite VIII far from
its margin; distance between the median posterior pair about 0.065—0.070 mm, to the
nearest hair 0.040—0.065 mm. No traces of wax glands visible on any part of the body.
Integumentum densely covered with mostly rather blunt nodules, on middle of
mesonotum with basal width of about 0.004 mm, at mutual distances of 0.007—0.010
mm; more caudad these nodules smaller, subdivided and more acute, in rows of 2—3;
on tergite VI acute and in rows of 2—4; on tergite VIII very small, in long wavy
transverse lines of 10—30 spinules, but larger and in short rows on the margins of that
segment. Head with a very broad median band of small nodules, and more inconspicuous,
scattered ones more laterally. Front with a large median bulge, which protrudes about as
much as the length of ant. segment II; this bulge not a marked process, in the middle
with 2 pairs of hairs of about 0.030—0.035 mm, one above the other, but sometimes
below the upper pair an extra pair of hairs half as long. Antennae 3/5—7/10 of length
of body, with segments I, II and basal 1/2—2/3 part of segment III pale, the rest dark
brown to black; flagellum faintly imbricated and imbrication more or less denticulate,
not spinulose on segments III—V, but on VI very indistinctly spinulose; in specimens
from Idaho with 0—1 rhinarium (5 segments with together 1 rhinarium), in specimens
from California with 1—3 always quite small and inconspicuous rhinaria on III, half-way
or past middle of segment; segment 14—18 times as long as its largest width, with
16—20 small hairs up to 2/5 of diameter at its constricted base. Rostrum unusually short,
not reaching mesosternum, last segment triangular with slightly convex sides, acute, only
1/3—3/8 of length of second joint of hind tarsi, with 2 hairs besides the 3 subapical
pairs. Legs faintly tinged; tibiae over about 34 of their length only somewhat spinulose
on dorsal surface and also near apex with rather few spinules; hind tibiae about 14 of

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Cau.
body a IV V VI on III

Il 2:62 1.54 0.50 027 0.27 0.16 + 0.17 1&2 0.12
2 2.90 1.88 0.57 0.37 0.34 0206-1019 1&1 0.12
3 2.47 1.68 0.55 0.31 029 0.18 + 0.18 17822 0.12
4 2.79 1.57 0.52 0.31 0.26 015 = 0.17 2.863 0.13
5) 2.68 1.58 0.52 0.30 0.27 0.16 + 0.17 1&1 ?

6 2599 1.54 0.51 0.29 0.26 0.16 + 0.16 0&0 0.12

(1—5, from Carex sp., U. C. Sagehen Creek (Sierra Co.), California, USAFLSVNEET:!
leg. D. Calvert no. C 67—7—18 a; 6, from “grass”, Cascade, Idaho, U.S.A., 18.VI.'54,
leg. G. F. Knowlton no. 495).

148 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

length of body, their apical hairs not differentiated; first tarsal joints normally spinulose,
with 5 ventral hairs; second tarsal joints of hind legs 14 of length of hind tibiae;
empodial hairs setaceous. Porus of siphunculi about 0.040 mm wide, hardly raised.
Tergite VIII in the shape of a rectangular triangle with hardly convex sides the top of
which is replaced by a circle with the length of the segment as radius; length 0.45 times
base; posterior margin evenly curved, not indented in the middle. Cauda with the knob
1.1 times as wide as long, rather evenly rounded, with 12—14 hairs.

Alate viviparous female.

Colour in life not known. In mounted specimen body about 2.15—2.50 mm long,
elongate; head laterally blackish brown with a pale median stripe and a pale brownish
area mesad of each eye; thorax unevenly dark, abdomen with large, dark brown marginal
sclerites and with broad, paler brownish spino-pleural cross-bars on each tergite; the bars
are narrower laterally, tend to fade or disappear in the middle on tergites I—III, are
fused with marginal sclerites only on tergites VI and VII, sometimes V, and may be
fragmentary on tergites I—III; tergite VIII evenly dark brown. Mesonotum with small,
very elongated nodules, but sclerites on abdomen only with long, wavy lines of small,
blunt spinules. Bulge on front lower than in apterae. Antennae 3/4—4/5 of length of
body, blackish, with the very base of segment III paler, with only processus terminalis
very indistinctly spinulose; segment III with 9—11 ciliate, larger and smaller rhinaria
more or less in one row, larger ones transversely oval, distal 1/6—1/5 part without
rhinaria. Fore wings long and narrow, with thick brown veins; sector radii very short,
sometimes hardly developed, very little curved; radial cell not or hardly wider than
stigma. Legs brown to dark brown, rather slender. Knob of cauda brownish black. Other
characters as in apterous viviparous females.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Cau.
body III IV V VI on III

1 2.15 1.56 0.52 0.33 0.27 0.14 + 0.16 9 & ? 0.12
2 2.47 1491 0.65 0.36 0.33 018-4407210 MIO Eil 78042
5) 2:30 1.88 0.64 0.36 0.33 0.19 + 0.18 10&10 O12

(1—3, from Carex sp., U. C. Sagehen Creek (Sierra Co.), California, U.S.A., 18.VII.’67,
leg. D. Calvert no. C 67—7—18 a).

Embryos. Marginal hairs spiny, from 0.016 mm on abd. segment III to 0.030 mm
on segment VII. Spinal and pleural hairs smaller, from 0.006, to 0.012 on tergite VII;
hairs on tergite VIII about 0.040 mm long.

Discussion. One sample of 5 good apterae and 4 alatae, one without head, is available
from California, and a sample of 4 much damaged apterae from Idaho. The samples
differ very little except in the number of rhinaria in the apterae. The California sample
was mixed with Thripsaphis verrucosa Gill, of which one aptera and one alata was
present. This made it possible to decide that the differences between the two highly
similar species were not caused bij eco-factors. But something else may cause confusion.
In the bodies of these aphids one may find larvae of hymenopterous parasites with a very

D. Hite Ris LAMBERS: On American aphids 149

warty skin similar to that of T. verrucosa, but with the warts more closely packed. Such
a larva in T. scabra might lead to misidentification. Then the different shape of abd.
tergite VIII or the much less reliable shape of the front could help.

Alatae of both species have more pronounced differences in frontal structures, but
also the sculpture on the sclerotic parts of the abdomen is very different. On tergites
II—V this consists pleurally in verrucosa of transversely oval warts with 2—4 blunt,
dark spinules on their posterior margin but in T. scabra there are only rows of 3—7
spinules.

It is doubtful whether the curious shape of the sector radii is a good discriminant. The
veins in this sample show often some extra forks, and therefore the shape of the sector
radii might be teratological.

Types. Holotype: apterous viviparous female (no. 1 of measurements), from Carex
sp., Univ. of Calif. Sagehen Creek (Sierra Co.), California, U.S.A., 18.VII.'67, leg.
D. Calvert, no. C 67—7—18 a. Paratypes: apterous and alate viviparous females with
collecting data as for holotype; apterae viviparae (damaged), from ‘grass’, Cascade,
Idaho, U.S.A., 18.V1.’54, leg. G. F. Knowlton.

Thripsaphis (Trichocallis) utahensis Knowlton & Hall, 1950

Richards (1971) classifies this species as an Allaphis Mordvilko, which in Richards’
sense is “without clusters of wax pores”. Through the kindness of Dr. Louise M. Russell
I could borrow the type slide from the U.S. National Museum, Washington, D.C. This
contains three remounted specimens: one adult apterous viviparous female, one alatoid
nymph, and one alate viviparous female. The aptera, which I choose as lectotype, and
the alatoid nymph have large fields of wax glands on abd. tergite VIII, around the
posterior spinal hairs and marginally on abdomen. A slide marked “Paratypes IV”, from:
grass sweeps, Logan, Utah, 15.V.’43, borrowed from the U.S. Nat. Museum, contains
two apterae viviparae in which the wax glands are also quite distinct although these
specimens are still in balsam.

Thripsaphis (Trichocallis) verrucosa Gillette, 1917

The published descriptions seem to cover three taxa which are treated as subspecies.
They key as follows:

1 (2) In apterae legs and antennae dark to blackish brown, and also body conspicuously
brown with a pale median stripe, and dark brown intersegmental transverse lines.
Tergum evenly covered with semiglobular or slightly transverse nodules, little
larger than the socket of a dorsal hair, but taller. Ant. segment III with rhinaria.
In alate mesonotal lobes with randomly arranged, nearly round nodules. On
Carex. Labrador and Alaska. T. verrucosa nodulosa subspec. nov.

2 (1) In apterae legs pale, antennae with at least segments I and II and a basal portion
of segment III pale. Ornamentation of tergum of a different kind. Ant. segment
III with or without rhinaria.

3 (4) In apterae ant. segment III without rhinaria in all available samples. Antennae
about half as long as body or shorter. Integumentum on mesonotum to abd.
tergite III sometimes with large, quite flat, transversely oval or eye-shaped warts

150

4 (3)

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

but mostly subdivided into 1—3 nodules on their posterior half, caudally and
medially soon splitting into pairs or triplets of small warts; undivided larger
warts at least in one direction farther apart than their own longest axis; on
margin of body warts rather acute and all inclined caudad. Sclerotic bars on
abdomen of alate ornamented with rows of tiny, acute spinules not on the
posterior margin of flat scales. S.W. California.
T. verrucosa subverrucosa subspec. nov.
In available material at least one ant. segment III with one or more rhinaria on
distal half. Antennae rarely half as long as body, usually considerably longer.
Integumentum from pronotum to about abd. tergite VI in the middle covered
with eye-shaped or strongly transversely oval, never subdivided, very flat warts,
which stand at most as far apart as their shortest axis, while their longest axis is
about as long as the dorsal hairs on these tergites. On more posterior tergites
spinules appear on the posterior margin of the warts. Alatae on mesonotal lobes
with oblique rows of strongly oval nodules, on the sclerotic bars of abdomen
with transverse oval warts bearing small spinules on posterior half (Photograph
10 b, p. 14 in Richards, 1971). All European material, and samples from
Colorado, Utah, California and Quebec. May occur mixed with the preceding
subspecies. (T°. leporinae Börner).
T. verrucosa verrucosa Gill.

Thripsaphis verrucosa nodulosa subsp. nov.

Apterous viviparous female.

Colour in life not known. In mounted specimens body 2.45—2.85 mm long, spindle-
shaped, with largest width at abd. segment I at about the middle of the body, con-
spicuously brown with a pale median stripe from head to tergite VI, VII or VIII,
with interrupted, pale, membraneous, intersegmental lines, bordered caudad by narrow,
dark brown lines across the abdomen. For dorsal ornamentation see key to subspecies.
Median depression on posterior margin of abd. tergite VIII distinct, but the scabrous
processes on both sides very low. In those specimens in which tergite VIII is slightly
elevated, distinct wax glands or cribriform discs visible at the level of the marginal hairs,
especially distinct in no. 4 in the table below. Antennae dark brown to black with only
the very base of segment III pale; segment III with many more rhinaria than in the main
species. Legs dark brown to brownish black, only seemingly not much spinulose. Other
characters more or less as in T. verrucosa sensu stricto.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Cau.
body III IV V VI on III
1 2.60 1.43 0.47 0.27 0.21 0.15°737043 5&7 0.15
2 2.63 1.47 0.46 0.28 0.23 0.16 + 0.15 6 & 6 0.15
5) 2.64 1.40 0.47 0.26 0.19 Os 17E ENOR 486 0.15
4 2.54 1.59 0.47 0.27 0.21 0155015 7&7 0.15
5 2.69 1867 0.57 0.33 0.29 0.16 + 0.15 6&7 0.15
6 2.78 1.66 0.56 0.30 0.29 0.18 + 0.15 5&6 0.15

D. Hire Ris LAMBERS: On American aphids 151

(1—4, “Emergent vegetation”, S. Carter Basin (Labrador), Canada, 5.VIII.'58, leg.
B.S.E.S., British Museum no. 1958— 490; 5—6, from Carex sp., Juneau, Alaska, U.S.A,
26.VII. 61, leg. F. C. Hottes no. 10).

Alate viviparous female.
Colour in life not known. Material in very poor condition, but clearly as dark as
apterae. Ant. segment III with 13—17 rhinaria.

Measurements in mm.

No. Length Ant. Ant. segments Rhin Cau.
body III IV V VI on III
1 2.90 ? 0:77 0.38 0.32 0.18 + ? 17T 0.16

(from Carex sp., Juneau, Alaska, U.S.A., 26.VII.'61, leg. F. C. Hottes no. 10).

Discussion. The two samples of the subspecies are very much alike, but differ con-
siderably from all other material of T. verrucosa. Both come from rather far north, and
they could be modifications by low temperature. Yet they grew up during the hottest part
of the year, and they are much darker than samples that grew up early at high altitude.
The Alaska sample, from July, has one apterous male. This suggests that production of
this sex is not induced by short day and low temperature in this subspecies. In European
and Colorado populations of the main species sexuals are produced in late autumn.

The discovery of wax glands on abd. tergite VIII in viviparae of this subspecies with
its dark tergites suggest that these glands may well be present in the other, pale sub-
species. Even in dark forms they are very inconspicuous because in ‘“good’’ preparations,
with a quite horizontal tergite VIII, they are in the plane of the optical axis.

Types. Holotype: apterous viviparous female (measurements no. 1), in “emergent
vegetation”, S. Carter Basin, Labrador, Canada, 5.VIII.'58, leg. B.S.E.S., British Museum
no. 1958—490; paratypes: apterous viviparous females with collecting data as for holo-
type; apterae viviparae, two damaged alate viviparae and one apterous male, from Carex
sp., Juneau, Alaska, U.S.A., 26.VII.’61, leg. F. C. Hottes no. 10. Holotype deposited
in British Museum (Nat. Hist.) where also most paratypes from the same sample are.

Thripsaphis verrucosa subverrucosa subsp. nov.

Apterous viviparous female.

Colour in life not known. Mounted specimens 1.90—2.50 mm long, elongate spindle-
shaped with largest width mostly at metanotum, faintly brownish-yellow with narrow,
light brown, pleural intersegmental transverse lines, marginally slightly darker because
of the nodules in profile. For dorsal ornamentation see key to subspecies. Median furrow
on posterior margin of tergite VIII not very distinct. Antennae with at least segments I,
II and basal half of III pale, the rest brownish black. Legs coloured like the body; tibiae
conspicuously spinulose till base, except on the caudal side of the base. Other characters
as in T. verrucosa sensu stricto.

152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Measurements in mm.

No. Length Ant. Ant. segments Cau.
body III IV V VI
1 2.44 1.24 0.38 0:23 0.20 O15 094 0.11
2 197 0.99 027 0.17 0.17 0.14 + 0.13 0.10
3 2.00 1.00 0.30 0.16 0.17 OS IONS 0.10
4 227 JETS 0.34 0.21 0.18 0:14 40:15 0.11
5 2.01 1.09 0.32 0.19 OT, 0.14 + 0.14 0.10
6 257 1513 0.33 0.20 017 OG 10213 0.11
7 2:30 Wali) 0.33 0.20 0.18 OLS 20213 0.11

8 2.45 1.04 0.32 0.17 0.16 01561012 OL

(All from Carex sp., California, U.S.A., 19.V.’61, leg. R. van den Bosch; 1—2, Wheeler
Springs (Ventura Co.), no. 61—V—19 r; 3—4 Matilija (Ventura Co.), no. 61—V—
19 s; 5—8, Gorman (Los Angeles Co.), no. 61 —V—19 t).

Alate viviparous female.

Colour in life not known. For ornamentation of abdomen see key. Lobes of mesonotum
not well visible but with oblique rows of rather dispersed spinules besides an indistinct
pattern of broad oblique striae. Other characters as in T. verrucosa sensu stricto.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Cau.
body III IV V VI on III

1 2:19 155 0.49 051 025 0.17 +10.17 8&11 0.11

(with apt. viv. fem. nos. 1—2).

Discussion. Dr. van den Bosch was after hymenopterous parasites of Carex aphids,
and in one trip collected in three places, probably on the same Carex. The Wheeler
Springs sample was a mixture of T. verrucosa sensu stricto, the subspecies described
above, a species described in this paper as T. hybrida, and one T. ossiannilssoni. The
other samples are quite homogeneous.

The absence of rhinaria in all the apterae made me think that perhaps the specimens
were fundatrices of T. verrucosa sensu stricto. That, however, is not so likely because
there was also an alate, which clearly belongs with the apterae.

The subspecies is not a typical California form of verrucosa in as much that also quite
normal verrucosa from several other localities, and even from Wheeler Springs, are
available.

Types. Holotype: apterous viviparous female (measurements no. 6) from Carex sp.,
3 miles S.E. of Gorman (Los Angeles Co.), California, U.S.A., 19.V.’61, leg. R. van den
Bosch no. 61—V—19 t. Paratypes: apterae viviparae with collecting data as for holo-
type; apterous viviparae and one alate viviparous female with collecting data as mentioned
under measurements 1—4.

D. Hire Ris LAMBERS: On American aphids 153
Uroleucon sijpkensi spec. nov.

Apterous viviparous female.

Colour in life brown, with black siphunculi, pale cauda, pale legs, and pale antennae
with dark apices. In mounted specimens body 2.80—3.35 mm long, broadly spindle-
shaped. Dorsal hairs stiff, with spear-shaped apices, spinally on abd. tergite III to 0.065
mm long, on tergite VIII slightly longer, nearly all placed on brown scleroites up to
0.050 mm in diameter; no trace of an antesiphuncular sclerite present, but postsiphuncular
sclerites well developed; tergite VIII with 4 hairs. Marginal tubercles on abd. tergites
II—IV mostly present, dark, and smaller and flatter than the socket of a dorsal hair,
hardly visible on a small sclerite between two marginal hairs. Head light brown, smooth.
Frontal tubercles strongly diverging; depth of frontal furrow just over 14 of distance
between antennal bases. Antennae considerably longer than body; segment I slightly
darker than head; segment II as dark as head; segment III with part bearing rhinaria
darker to very much darker than head, basal and apical part conspicuously paler, the very
apex dark brown to blackish, with 17—36 faintly protruding rhinaria of very variable
size, with bulging membranes, mainly along one side over basal 4/7—2/3 part of
segment; segment IV basally about as pale or dark as distal part of segment III, apicad
darker, to brownish black like the rest of flagellum; proc. terminalis longer than segment
III, 514—6 times as long as basal part of segment VI; longest hairs on segment III
slightly longer than basal diameter of segment. Rostrum (stylets) reaching to hind
coxae; last segment about 0.18—0.20 mm long, about 1 1/3 times second joint of hind
tarsi, with 7—10 accessory hairs. Legs evenly yellowish, to slightly brownish yellow, with
the femora dorso-apically on distal 1/4—1/5 (hind legs) to distal 1/3 (fore legs)
suddenly brown to blackish brown, tibiae similarly dark at the articulation, and on distal
14, part; first tarsal joints with 5 or often, on some legs, with 4 hairs. Siphunculi evenly
black, rather variable in length, from 2/9—3/11 of length of body, tapering from the
thick base, with distal 1/3—1/4 part cylindrical and there 1 1/5 times as thick as mid
portion of hind tibiae, with distal 1/4—2/7 reticulated, the rest to the very base slightly
and inconspicuously imbricated. Cauda pale, ensiform and nearly 3 times as long as width
at its very base, slightly blunt, with 12—18 hairs of two conspicuously different sizes,
all with fine apices.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Siph. Cau.
body at IV V VI on III
2 OOM 095 ND CO 059. 704974 1.06, 20821 0770 10593
2 a 3.69 0.93 0.71 0.59 0.18 + 1.04 30&34 0.88 0.57
3 3.06 353 0.86 0.65 0.55 0218 + 1:05 19.8219) 0:68 0.51
4 3.26 3.66 0.91 0.73 0.57 0.17. + 1.02 30,836 0:87 0.55
à 2.89 3.58 0.87 0.68 0.57 0.17 + 1.03 25&26 0.68 0.47
6 3.13 3.58 0.88 0.68 0.57 0.18 + 1.04 21&22 0.71 0.48
7 3.14 a) 0.86 0.66 0.55 OS 1.05 18220772 0.53
8 ZI 3.40 0.85 0.63 0.54 OLS O7 SO 0570 0.47

(1—8, from Solidago macrophylla, Gaspé Nat. Park, Quebec, Canada, 16.VIII.'65, leg.
J. P. Sijpkens no. 94).

154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 4, 1974

Oviparous female.

Like apterous viviparous female, but abd. tergite VIII with up to 8 hairs; hind tibiae
quite yellow with short black socks, considerably swollen to up to 214 times maximum
width of other tibiae, with a great many pseudosensoria. Siphunculi thinner, from a thick
base rapidly tapering to the cylindrical distal 2/3—3/4 part, rather scabrously imbricated
from base to reticulated area. Cauda much shorter and also thicker.

Measurements in mm.

No. Length Ant. Ant. segments Rhin. Siph. Cau.
body III IV V VI on III
1 3.04 9 0.96 0.80 0.64 0.19 Ete 29&31 0.80 0.45

2 gal 3.66 0.92 0.75 0.66 0:20,:530.87; 5, 20E En 0199, 0.45
(1—2, with the apterae viviparae).

Discussion. The principal character of this species is the paleness of the legs. In quite
a number of brown North American Uroleucon (formerly Dactynotus) the tibiae are
partly pale and transparent on the basal half, at least in apterae viviparae. But then the
hind tibiae in oviparae (as far as known) are evenly black. In U. sijpkensi the hind tibiae
are yellow also in oviparae with only a small part at the apex blackish.

Dr. A. T. Olive, to whom I sent a slide, identified this as U. nigrotuberculatum
(Olive). Biometrically it agrees with that species, also in the tiny dark marginal tubercles,
but not in the pigmentation of the flagellum and of the hind tibiae which are blackish
in paratypes, in other material identified by Dr. Olive, and in several Canadian samples.
The siphunculi in viviparae of U. nigrotuberculatum are, with the exception of few often
invisible imbrications on the underside very near the base, completely smooth to near the
reticulated area. In U. sijpkensi they are very lightly imbricated in apterae viviparae, but
conspicuously imbricated in oviparae.

Two apterae from Solidago sp. in Canada were sent to Dr. W. R. Richards in Ottawa
who confirmed my suspicion that they were his U. pielozi. The apterae in that sample
have a considerable part of the tibiae pale, but about halfway they become dark towards
the apex. The oviparae have evenly black hind tibiae. The flagellum is black with only
the very base of segment III paler. I have not succeeded in finding a difference between
pieloui and the earlier described U. hieracicola (H.R.L.), of which pielout may well be
a synonym.

U. sonchellum (Monell) strongly resembles U. sijpkensi spec. nov. in the pigmentation
of tibiae, but the femora are more extensively black at the tips, and the short siphunculi,
and the extremely long ant. segment III with its many rhinaria exclude confusion with
U. sijpkensi.

According to Dr. Sijpkens the aphids infested the inflorescences of their host. It is
remarkable that oviparae, one teneral, the other old, were collected as early as the middle
of August, together with the apterae viviparae.

The species is named after my compatriot and friend, the late Dr. J. P. Sijpkens, who
made very carefully documented collections of aphids in many parts of the world.

Types. Holotype: apterous viviparous female (measurements no. 6, in one slide with
an ovipara), from Solidago macrophylla, Gaspé Nat. Park, Quebec, Canada, 16.VIII.’65,
leg. J. P. Sijpkens no. 94. Paratypes: apterous viviparous females, larvae and oviparae
with collecting data as for holotype. Some paratypes in the collection of Dr. A. T. Olive,
Winston-Salem, North Carolina, U.S.A.

D. Hire Ris LAMBERS: On American aphids 155
TYPES

Unless otherwisely stated, types of new species and subspecies described in this paper
are in the author’s collection.

ACKNOWLEDGEMENTS

The ready cooperation of the following colleagues, who sent valuable material as
gifts or on loan, or established the identity, of species mentioned in this paper is
gratefully acknowledged: Drs. R. van den Bosch; Maria P. Bozhko; R. C. Dickson;
V. F. Eastop; W. D. Fronk; G. F. Knowlton; the late C. F. Lagace; A. T. Olive; Miriam
A. Palmer; W. R. Richards; Louise M. Russell; the late J.P. Sijpkens; H. L.G. Stroyan;
D. J. Voegtlin; H. Weidner; and especially G. Ch. Shaposhnikov who refound and made
available Mordvilko material of crucial importance.

Mrs. M. Hielkema-Visser made the drawings in this paper.

REFERENCES

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Eastop, V. F., 1966. — A taxonomic study of Australian Aphidoidea (Homoptera). — Austral. Jl.
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Gillette, C. P., 1917. — Two new aphid genera and some new species. — Can. Ent. 49: 193—199.

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Biol. Zentralanst. 78: 1—52.

Richards, W. R., 1971. — A synopsis of the world fauna of the Saltusaphidinae or sedge aphids
(Homoptera: Aphididae). — Mem. Ent. Soc. Canada 80: 1—97.

Stroyan, H. L. G., 1960. — Three new subspecies of aphids from Iceland (Hem., Hom.). — Entom.
Medd. 29: 250—265.

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Daten IAP MUS. COMP. ZOOL .
ah Sl. ok ; LIBRARY
huy 18 1974
HARVARD
AFLEVERING 5 UNIVERSITY 1974

= TIJDSCHRIFT
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| Habrophorula, a new genus from China (Hymenoptera, Anthophoridae), Ps 157—
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ok die id SE

EEE a ee ee Tl ac er alie zee lings a nn

REVIEW OF CENTRAL AND EAST ASIATIC HABROPODA
F. SMITH, WITH HABROPHORULA, A NEW GENUS
FROM CHINA (HYMENOPTERA, ANTHOPHORIDAE)

M. A. LIEFTINCK
“Kalliste’, Rhenen U., Netherlands

With 1 table and 100 text-figures

CONTENTS
Introduction . . >». » ees Pe inne
Census of material and Liana REEN o NA es
Senetalscemarksion classification and relationships …. . . en At 159
Isrographicaltandiseasonal.distribution. ge We , . |. tt ae et et
IDeseriptiverkeys and ıllustrations i Rei. ee te 161
BRECREORERERSDEGIESMIALES ERE Sw we en ee ROS
Key to the species, females i Sick IE Des, serie i ARE FE MM EET
Discussion of species and subspecies. ME 1 eH LY
H. pekinensis Ckll. — ® N China, redefined: 4 al reden Beel yt, Moe UO
H. sinensis Alfken — & Q E China, types and further material redefined . . . . . 180
Er zumidınons spec. nov. — 9 South VietNam’. . -. … von « … . 182
H. orbifrons spec. nov. — 9 Thailand. . oe ee. ei on 115
H. mimetica Ckll. — & 2 SE China, redefined; ssi di JT OR ORAN LS SEES
messcanota spec. nov.— 9. North VietNam) . renee er 2 7, te 2185
H. turneri Ckll. — & 9 Assam, redefined; first d . . LdC el 85
H. rowlandi Meade-Waldo — & 9 Assam, types and raten REA ne Se, ALS
BNN tainanicoia Strand — & 9 Taiwan, redefined; first 4. . . . . 191
EON ellassubspec. nov. — & 9 SE China oe 0,1 A 193
H. apostasia spec. nov. — & Nepal . PORN Er A06
H. sutepensis Ckll. — 3 9 Thailand, na Holotype 4 tE fy eene er. tu
Hikoate:Ckll. — 9 N India, topotype.redetined. .. . = Lu... ge 199
H. bucconis (Friese) — & 2 Taiwan, redefined; lectotype 6 . . . . . . . . 201
H. imitatrix spec. nov. — & 9 SE China . . DE 05,
H. radoszkowskii (D.T.) — & 9 N India and Nepal, redefined lectotype 4 DNS NTI 205
H. plantifera spec. nov. — & Assam . . . MNT at. 06208
H. pelmata spec. nov. — & Nepal. . ADEN pat Paes en a ne PO)
H. deiopea (Cameron) — 3 2 N India, redefined: ona 8. hr ee rr 212
H. apatelia spec. nov. — & 9 N India and 4 2 doubtful from Ne OEE staten 2115
H. krishna Bingham — & Sikkim. . 216
Habrophorula gen. nov., from Habropoda foi H. Pini Ckll. — è 9 SE Chine. redefined 217
References dg - ns ‘ 2 ; : sige 224
Gia, OMA EEE

157

158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974
INTRODUCTION

The object of the present paper is to round off, as far as possible, my investigations
of the general morphology and specific characters of the anthophorine genus Habropoda,
the emphasis being laid on the members inhabiting Central and East Asia. In a former
paper (Lieftinck, 1966) I have treated this genus in a more general way in connection
with allied genera, pointing out and illustrating the characters of the type species, Habro-
poda tarsata (Spin.) and five of its immediate allies which occur in the western part of
its range, i.e. the Mediterranean region and western Asia. A sixth species of the same
group, H. pekinensis Ckll. from East Asia, which I had not yet seen, is included in the
present article.

In my previous account brief comments were already given on the remaining taxa, ten
in number and all from Asia, but as these had not been studied in any detail, all were
left as species requiring further investigation. It must be said at the outset, that of the
species just mentioned one, uubilipennis Cockerell, originally described from a single
female, deviates so much from all others that it has to be segregated and redefined as the
first member of a distinct new genus, Habrophorula. The re-characterization of this bee
is based mainly on the hitherto unknown male, whose structures are fully described and
illustrated at the end of this paper.

CENSUS OF MATERIAL AND ACKNOWLEDGEMENTS

This study is based on the examination of many hundred specimens pertaining to 22
taxa, including nine which are new and one previously described species mentioned
above, which requires a new generic name. Besides these, redescriptions and figures are
given of 12 earlier named forms (including the types of five), the generic status of only
one still remaining uncertain.

Most of the material originates from the province of Fukien, in south-east China, and
the island of Taiwan‘). Thanks to the activities of Dr. T. C. Maa, who investigated the
insect fauna of several Chinese provinces for about seven years in succession, no fewer
than eight species of Habropoda and the undescribed male of nubilipennts Ckll., the type
species of Habrophorula gen. nov., were collected by him in China, Taiwan and Thailand.
Other important material was accumulated by members of the British Museum East Nepal
Expedition 1961—62.

Below the names are listed of the institutions and private collections from which
valuable material was obtained for the purpose of this study, preceded by the usual
symbols denoting the ultimate location of specimens borrowed. These data are followed
by the names of persons who generously gave me access to the collections under their
care and who provided many other facilities. To all these I wish to extend my grateful
thanks for the loan of types and other indispensable material necessary for the preparation
of the present article. I am particularly thankful to my colleague T. C. Maa, who
supplied important information on the biology of certain Chinese species and details
concerning the localities and distribution of these. Paratypes and duplicate specimens have
been distributed as much as possible among the institutions mentioned.

1) For details concerning H. Sauter’s localities in the island of Taiwan (Formosa), sec p. 201—202.

M. A. LIEFTINCK: Review of Asiatic Habropoda 159

BISH — B.P. Bishop Museum, Honolulu (J. L. Gressitt)

BM — British Museum (Natural History), London (R. W. Crosskey, I. H. H.
Yarrow and G. R. Else)

ML — Rijksmuseum van Natuurlijke Historie, Leiden

MNB — Museum für Naturkunde an der Humboldt Universität, Berlin (E. Königs-
mann)

NRS — Naturhistoriska Riksmuseum, Stockholm (S. Erlandsson)

OUM — University Museum, Hope Department of Zoology, Oxford (C. O'Toole)

SMF — Forschungsinstitut Senckenberg, Natur-Museum, Frankfurt a.M. (D. S.
Peters)

TCM — Tsing-chao Maa collection, Neihu

UKL — Department of Entomology, University of Kansas, Lawrence (C. D. Mich-
ener) i

USNM — National Museum of Natural History, Washington, D.C. (P. D. Hurd, Jr.
and K. V. Krombein)

ZIL — Zoological Institute, Acad. Sci. USSR, Leningrad (M. N. Nikolskaja and
V.I. Tobias)

GENERAL REMARKS ON CLASSIFICATION AND RELATIONSHIPS

The main problem that confronted me was that of the natural relationships of the
species involved and the question as to whether even Habropoda — after the segregation
of Elaphropoda (see Lieftinck, 1966) and Habrophorula (of this paper) — should be
further subdivided. It was suggested earlier that some at least of those left in Habropoda
could be linked to Emphoropsis, a nearly allied genus occurring in the Western
Hemisphere. It will be seen that further investigations proved the impossibility of doing
so and that a definite generic or subgeneric assignment of the group must still be left
in abeyance.

As with Anthophora, one of the difficulties in defining sections or subgenera is the
fact that practically all characters available to establish these are unisexual and found in
the hidden sternal plates and genital organs of the male. In studying these, much was
learned of the various forms of their structure; giving particular attention to the internal
sternites, for example, these proved to be surprisingly differentiated, most species ex-
hibiting peculiarities and specializations of their own. It goes without saying that an
attempt was made to correlate these varying structures with variations in other organs,
such as the antennal and leg structure, the shape of the facial sclerites with their colour
marks, and the nature and colour of the vestiture. However, these features proved to be
allotted differently, so much so in fact that they greatly impeded the separation of well-
defined groups. Although it was fairly easy to pick out pairs, or even minor sections of
more closely associated species, there were too many strays that seemed to stand apart
from the others. Taking all characters together, and seeing that so many of them are
shared out indiscriminately, it soon became evident that it was impossible to arrange them
in a harmonious system.

Turning our attention again to the New World genus Emphoropsis. Though some of
the Asiatic species were temporarily attributed to this genus by Michener (1944) and
myself, I must confess that my own suggestions to that effect were rather premature. In
point of fact nearly all taxonomic characters of the present composite section of Habro-

160 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

poda seem to meet (and intergrade with) those of Emphoropsis, which obviously
constitutes an equally diversified group. The existing definitions of Emphoropsis are
meagre, Cockerell’s review (1905) being not exactly “according to Cocker”, neither do
they do justice to the interspecific variation observed in this genus (see in particular
E. excellens Timberlake, 1962). Hence it is now Emphoropsis which is in need of a
revised definition as a genus, while its members require a thorough analysis of their finer
structures. Unfortunately, only the type species, E. laboriosa (F.), appears to have been
investigated upon the male copulatory organs. True, the genital capsule with its ap-
pendages does not reveal differences of any great importance as compared with that seen
in Habropoda. Nevertheless the chief claim to generic rank of Emphoropsis (i.e. the type
species) would appear to be the complete absence of a pair of heavily scleroid transverse
ridges at the base of the disks of the seventh and eighth gastral sternites. These strong
raised transitional carinae are conspicuously present in practically all Habropoda, inclusive
of the species presently discussed. On balance, for the present this peculiarity supplies the
strongest motive to leave the genus as it now stands, with all Old World members
included. Future investigations on a broader scale and on the basis of a great many
characters may necessitate subgeneric divisions of both Habropoda and Emphoropsis, and
it is not unlikely that the taxa here treated will have to participate in this.

GEOGRAPHICAL AND SEASONAL DISTRIBUTION

In the accompanying Table I have summarized the available data relating to the
geographical distribution of the species here treated, with as many particulars on seasonal
and altitudinal occurrences as could be accumulated from the literature, the locality labels
of museum specimens, and from information supplied to me by field workers.

It seems that throughout the Himalayan range the most favourable reproduction period
for these bees is from August to October, as it also appears to be in the mountains of
Shillong, in Assam. Similarly, at the higher levels of Nepal, most Habropoda were
collected just after the summer rains, which characterize the monsoon season, i.e. in the
period between July and November, the monsoons in Nepal occurring regularly between
June and September. In this connection the re-appearance of H. radoszkowskit again in
April and May, is hard to explain, two generations being not likely to be effected at
these high altitudes.

Interesting differences in the flight seasons can be observed in some of the Chinese
Habropoda occurring south of the Yangtze River (about lat. 30° N). One of these, H.
sinensis, is widely distributed in the submontane countries; but whereas this, and four
others in south-east China and Viet Nam, were all assembled in a period between June
and late autumn, the somewhat isolated faznanicola, from Taiwan, as well as its subspecies
maiella from the opposite continent, appeared much earlier in the year, viz. from March
till May. Morphologically this taxon stands rather apart from the others, which may or
may not have something to do with the fact that both races afford good examples of
early spring bees restricted to hill-country at lower altitudes. Dr. T. C. Maa regularly
kept me informed about his efforts in Taiwan to find /ainanicola during the autumn
months, from August onward, but he never once came across a single individual until
the emergence of the next spring generation. For notes on the nesting habits, see under
that species. A second Taiwanese member of the group, bucconis, was recorded by other
collectors as having been captured from October to January, but this species was not yet

M. A. LIEFTINCK: Review of Asiatic Habropoda 161

TABLE I. REGIONAL AND SEASONAL DISTRIBUTION OF CENTRAL AND EAST ASIATIC Habropoda ET AL.

Name Region Altitude Months of the year Un-
(W to E) (in metres) 1 2 3 4 5 6 7 8 9 10 11 12 known

hookeri W Himalaya 2300 AR gr Ap

deiopea W Himalaya 2300— 2500 + +

apostasia Nepal 2000 ap an

pelmata Nepal 2000 ie

krishna E Himalaya 2300 +

apatelia Nepal; E 2000—2300 al
Himalaya

radoszkowskii Nepal; E 1465— 2450 + + qe an on
Himal.; Assam

plantifera Assam 1400—1500 +

turneri Assam 1460 Te or

rowlandi Assam 1460 SP SE È

sutepensis Thailand 1650 ca.

orbifrons Thailand 1500

tumidifrons S Viet Nam 1400—1500

disconota N Viet Nam low level

sinensis SE China 350—2000 +++ + +

mimetica SE China 300—500 sia

imitatrix SE China 500—1500 ar SP Ar

t. maiella SE China 400—1000 dE op

t. tainanicola Taiwan 250 +++

bucconis Taiwan 0—500 ca. + + +

RN SG a DO DOM TT

Habrophorula SE China 1000 +

nubilipennis )

encountered by Dr. Maa, who suggests that it may occur in other parts of the island in a
different season. It is, perhaps, significant that imitatrix, the nearest ally of bucconis from

continental China, was found abundantly each year by the same collector from August
to November.

DESCRIPTIVE KEYS AND ILLUSTRATIONS

Having in mind an adequate valuation of characters, I tried to construct suitable keys
for both sexes, whereby I had to choose between two possibilities, (1) to identify species
merely on some characters apt to vary (such as the presence or absence of yellow clypeal

162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

marks, or a spot on the antennal scape); or (2) by trying to give an impression of the
whole insect by sorting out the various taxa in a descriptive key by a combination of as
many salient particulars as could be detected. The second course has been followed as the
most satisfactory one: first to prevent misidentifications liable to happen when too few
and superficial distinctives are used, and further, to avoid unnecessary repetitions in the
specific descriptions. Females of closely allied species being notorious by their homo-
geneity, the characters will be found to differ widely from those given for the males.
The difficulties engaged with defining clear-cut group characters for the males are fully
expressed in the keys and especially brought out by the illustrations. These only partly
reflect true relationships, but I do hope that they show how the characters are shared by
each. The sequence in the text that follows will be found more in accordance with a
certain phylogenetic continuity. Additional information on peculiarities of structure is
given under each species.

Concerning the illustrative methods followed it can be said that all illustrations are
original camera lucida drawings, except Fig. 7—10, of the face markings of some species,
which are semi-diagrammatic and on the same scale. The hind legs are shown in full
profile view; though most of the pile covering the tibio-tarsal surfaces is omitted, it was
necessary for each species to indicate clearly the contour of the strong marginal hairs of
these segments, so as to show the length and density of the vestiture: a very important aid
in species recognition. The compact bristly hairs covering and fringing the exposed part of
the apical tergite of the males are omitted from all figures. As customary, the separate
parts of the invaginated sclerites and genital organs were dissected out, cleaned, and
depicted from glycerin slide preparations under cover glass, hollow slides having been
used for the genital capsule to prevent pressure and to keep the ventral surfaces in
horizontal position. For ultimate preservation all terminalia were transferred, without
excess glycerin, to cellophane capsules glued to a card on the pin with the specimen. It
should be noted that corresponding organs have not in every instance been drawn on the
same scale.

KEY TO THE SPECIES, MALES

N.B. — The males of H. disconota, hookeri, orbifrons and tumidifrons are still unknown

ile Legs robust and strongly modified: fore coxae armed with a long process, fore
tibiae with an apical tooth, and conspicuous tooth-like projections also at the inside
of the first two protarsal segments; hind basitarsus with great flattened antero-apical
lamina. Mid and hind femora posteriorly with distinct longitudinal keel. Tarsal
arolia well developed. Ocelli arranged in a broad triangle; interocellar (IOD) and
ocellorbital (OOD) distances equal (hence IOD : OOD = 100 : 100); ocelloc-
cipital distance not longer than diameter of nearest ocellus. Labrum dark, only
slightly broader than long, its apex a little protuberant; disk clothed thickly with
long tuft of decumbent pale hairs directed anterad. Face, including paraclypeal areas
and a band along epistomal suture on supraclypeal area, bright chrome; clypeus not
unusually broad and bare, its surface convex, but not very shining, superficially
finely punctate; colour bright chrome, with pair of black marks, constricted at
middle. Antenna of great length, scape naked, anterior face broad, flattened and
conspicuously yellow; flagellum black. Eyes convergent below, inner orbits nearly
straight. Light dorsal pubescence of thorax long and dense, unicolorous. Marginal
cell of fore wing a little shorter than distance from its apex to wing tip. Integument

M. A. LIEFTINCK: Review of Asiatic Habropoda 163

of gastral tergites 1—6 black, evenly closely punctate; 1 clothed abundantly with
long, erect light hairs, which become shorter, decumbent, more condensed and band-
like apicad; most of 2—6 with mixture of erect black hairs varying in length, but
apical, margins of all with complete, sharply contrasting light bands made up of
compact, appressed hairs entirely concealing surface. Apical margins of sternites
lacking continuous hair-bands, but sides of 2—6 with long pale fringes. Tergite 7
broad, flattened dorsally, apex narrower and rounded. Apical sternal plates and
genital organs not studied. See description. Body a 15.0 mm approx. Hab.: W
ANNE China. ni Ee pekinensis

Legs more slender: ae Lx mail o not {ich keeled posteriorly,
only apical portion of hind basitarsus occasionally expanded and/or modified.
Remaining characters combined not as above . . 2

Integument of tergites 1—6 brownish black or Blade ena hema sea
closely and deeply punctate, punctures equal to or larger than interspaces; 1 clothed
with longish, raised light hairs, which become depressed, more condensed and
band-like apicad; most of 2—6 covered sparsely with short black hairs, but apical
margins of all with complete, rather broad and compact, sharply contrasting, light
ochraceous-buff to pale orange-yellow bands occupying at least one-fifth of the
exposed surface of each, these bands composed of thick, appressed and finely
branched, curled hairs entirely hiding the surface. Free border of sternites 2—5
straight, surfaces dark brown, margins narrowly paler, each with continuous fringe
of rather long yellowish white hairs of almost equal length, less compact than on
tergites. Apex of sternite 6 obtuse-angulate, with dense fringe of short pale marginal
hairs. Labrum dark, almost two times as long ‘as broad, widest basally; anterior
border gently convex, with tiny median emargination, a pale hair tuft in the depth
of incision, and a pair of minute subapical tubercles at the end of a low arched ridge
that runs almost parallel to apical margin; disk flat, closely, coarsely and rather
deeply punctate; pubescence short, pale, not concealing surface. Face broad, clypeus
convex but only little protuberant, almost totally hairless, its surface smooth and
shining, covered with large, widely spaced, rather superficial punctures, except
basally, where punctures are smaller and crowded; colour mainly bright chrome,
only sides with dark stripe along paraclypeal suture, widest and somewhat angulated
at level of obscured tentorial pits. Mandible bases and a low, triangular or trape-
zoidal stripe, tapering at either end, bordering the almost straight epistomal suture
on supraclypeal area, likewise bright chrome; paraclypeal areas unmarked, with
short appressed silvery hairs (Fig. 7). Malar space distinct, but narrower than
pedicel of antenna. Antenna slender, scape slightly curved, long and cylindrical, uni-
form red- to dark brown, anterior face clothed with long hairs; 3 much shorter than
4 + 5, ratio of length to width 100 : 70.4; length ratios of 3, 4 and 5 = 100 :
86.5 : 90. Ocelli forming an almost equilateral triangle; IOD : OOD = 70 : 100;
ocelloccipital distance very short, subequal to diameter of nearest ocellus. Eyes
parallel or nearly so, inner orbits somewhat convex mesially and a little diverging at
either end. Legs slender, pubescence rather short, hind tibia and basitarsus, Fig. 7;
tarsal arolia well developed. Marginal cell of fore wing longer than distance from
its apex to wing tip (ratio 100 : 74.6). Pubescence in front of head and dorsum
of thorax ochraceous to brownish yellow, rather dense though not very long; hairs
finely plumose and more distinctly brown at sides of face, upon vertex and occipital

164

34:

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

area; laterally all hairs palest yellow on genal area and thoracic sides. Tergite 7
closely punctate, apex slightly upturned, truncated, whole surface black-haired.
Sternites 7—8 and genitalia, Fig. 2—3; 7 strongly constricted about halfway length,
at which level transverse ridges are non-apparent and hardly sclerotised, apical
ee diamond-shaped (Fig. 2). Body oe 11.0—12.0 mm. Hab.: E China
sinensis

sii of (csi, ellen hone to black Be: al margins invariably
broadly pale, frequently semitransparent, thus accentuating the light hair bands,
when present; if so, then the latter are less sharply defined anteriorly: basal portions
of 1—6 usually at least with admixture of much longer, partly raised hairs, hence
never very short and quite black; if the first three or more segments are dark-
haired, then there are no well defined, compact, light apical bands . . . . 3
Eyes very large and bulging: shortest interorbital distance (across antennal
sockets) only about one-half length of eye; inner orbits in frontal view subparallel
but distinctly convex mesially, diverging gradually at either end. Labrum much
broader than long (about 100 : 60), disk flat, shining, coarsely rugosely punctate,
with smooth median line or low ridge, and clothed sparsely with short pubescence;
anterior border with broad and deep crescentic emargination carrying a tuft of stiff
pale hairs in the depth of the incision (Fig. 16). Ocelli in almost equilateral
triangle; IOD : OOD = 71.4 : 100. Head, including mandibles, labrum and
paraclypeal areas, black; clypeus entirely chrome, except a narrow lateral black stripe,
equal in width to the narrow paraclypeal area at its upper end; also a transverse
chrome bar, broadest at middle, along the slightly arched epistomal suture, on
supraclypeal area. Malar space nil. Antennal scape black; ratio of length to width
of segment 3 = 100 : 35.5; length ratios of 3, 4 and 5 = 100 : 35.5 : 44.4, Legs
brownish black, unmodified; hind basitarsus long, slender, laterally compressed,
slightly curved, parallel-sided, lacking a naked area on lower surface; coxae and
femora with mixture of light and dark pubescence, hind tibia and basitarsus clothed
throughout with rather long and dense black hairs, the dorsal fringe at hind
basitarsus longer than greatest width of the latter (Fig. 4). Tarsal arolia vestigial.
Marginal cell of fore wing longer than distance from its apex to wing tip (ratio
almost 4:3). Thorax and abdomen thickly and evenly clothed with rather short,
erect pale pubescence, without admixture of black, except a broad, ill-defined,
transverse dark band between the fore wings; pale hair on remainder of thorax and
tergite 1 light yellow (straw yellow, Ridgway), shorter, decumbent and deeper in
tint (ochraceous-buff) on 2—5, on which it forms very compact bands, ill-defined
anteriorly; tergites 6—7 black-haired. Apex of tergite 7 truncated. Sternites 2—5
unmodified, straight-bordered, each with apical fringe of long, wavy, . partly
recurved and mostly pure white, hairs, longest at sides; 6 bluntly triangular, with
dense fringe of short stiff marginal hairs; 7—8 and genitalia, Fig. 5—6. Sternite
7 constricted beyond halfway length, at which level the lateral ridges are small;
distal portion greatly expanded, but apex abruptly narrowed and ending in a slender,
rod-like median process. Body si 14.0—15.0 mm. Hab.: SE China (Fukien)
; mimetica
Byes not dl, legal is pai aad more E. separated above:
shortest interorbital distance always greater than one-half length of eye. Other
characters combined not as above. Generally smaller-sized species. . . . . 4

M. A. LIEFTINCK: Review of Asiatic Habropoda 165

Mid and hind tibiae and basitarsi black-haired, or almost so. Dorsal pubescent
pattern varicoloured: summit of head, anterior portion of thorax to behind tegulae,
as well as ends of tergite 2 and most of 3, black; remaining parts of thorax, tergite
1 and proximal portion of 2, canary yellow; distal portion of 3 and all succeeding
tergites orange to orange-rufous, the broad pale brown hind margins of 3—7
shining through the pubescence. Long silky hairs on genal area, sides and ventral
parts of thorax, the femora, and those fringing apical margins of sternites, almost
pure white. Labrum dark brown, much broader than long (100 : 75), its anterior
border evenly convex, apex slightly impressed medially but not excised; surface flat,
coarsely rugosely punctate, disk bounded by a low, wide, arc-like ridge running
subparallel to anterior border. Mandibles yellowish brown. Malar area black,
narrower than pedicel of antenna. Clypeus and paraclypeal areas almost entirely
bright yellow, their surface superficially punctate; also a narrow, transverse bar of
yellow, widest at middle, along the straight epistomal suture, on supraclypeal area;
clypeus marked with black laterally much as in sirensis, but black stripe gradually
broadened below. Inner orbits subparallel. IOD : OOD = 66 : 100. Ocelli in
isosceles triangle. Antennal scape yellow anteriorly and clothed densely with long,
soft, white pubescence; ratio of length to width of segment 3 = 100 : 42.5; length
ratios of 3, 4 and 5 = 100 : 52.5 : 57.5. Legs dark brown; hind basitarsus simple,
slightly curved but parallel-sided, much longer than broad (4 : 1); outer face
sulcate for its entire length, but no naked area on lower surface; hair fringing upper
and lower ridges shorter than greatest width of basitarsus (Fig. 17). Tarsal arolia
present, though of minute size. Marginal cell of fore wing longer than distance
from its apex to wing tip (ratio 100 : 81). Apex of tergite 7 shaped as in Fig. 18.
Sternites 2—5 straight-bordered, marginal hair-fringes more condensed and silky,
longest at sides of 3—5, whole surface of 6 clothed with short and dense, pale
rufous hairs. Sternites 7—8 and «Sa Fig. 19—20. Body length 13.0 mm
approx. Hab.: Assam . . “oiturnert

Mid and hind tibiae and tarsi at east cah light haie) oral pubescent body
pattern more uniform, never tricoloured. Tarsal arolia well developed and con-
spicuous . . Ba ED

Pubescence uniform buff alom, Beep that some 1. anca] mais upon head
and thorax are tipped with brown, and that there is an ill-defined but conspicuously
blackish brown transverse band of long raised hairs upon middle of tergite 2; sides
of segments clothed with still longer, finely branched light hairs. Labrum dark
brown, much broader than long (100 : 80), its anterior border a little projecting
and upturned, slightly crenulated medially, but not emarginate; disk somewhat
concave, not bounded by a ridge, coarsely rugosely punctate, well visible under the
long pale pubescence. Mandibles obscurely brownish yellow at middle. Malar space
black, somewhat broader than pedicel of antenna. Clypeus and paraclypeal areas
almost entirely pale yellow, a black paraclypeal stripe only bordering inner orbits,
and a minute brownish dot, or streak, to the inside of the obscured tentorial pits;
surface superficially punctate; also a pale yellow stripe, usually interrupted at middle
and occasionally absent, along epistomal suture on supraclypeal area. Inner orbits
subparallel but distinctly convex medially and diverging gradually below. IOD :
OOD = 80 : 100. Ocelli in isosceles triangle. Antennal scape dark brown, un-
marked with yellow in front, and clothed with long pale hair; ratio of length to

166

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

width of segment 3 = 100 : 51; length ratios of 3, 4 and 5 = 100 : 60.6 : 72.7.
Hind basitarsus simple, scarcely curved and subparallel-sided, much longer than
broad (3 : 1); outer face slightly concave, not sulcate, and no naked area on lower
surface; dorsal ridge with thin fringe of raised hairs, which are almost as long as
greatest width of basitarsus, especially the basal ones, those along ventral ridge
much shorter (Fig. 22). Marginal cell of fore wing distinctly longer than distance
from its apex to wing tip (100 : 71 approx.). Long silky pubescence on genal area,
sides and ventral parts of thorax, the femora, and at anteapical margin of sternites,
pale yellow. Apex of tergite 7 truncated, but hind margin with shallow crescentic
emargination (Fig. 23). Sternites 2—5 straight-bordered, marginal hairs at sides
much shorter and less compact than in /wrneri, especially at middle; 6 slightly
projecting, whole surface clothed densely with short rufous hairs. Sternites 7—8
and genitalia, Fig. 24—26. Body length 11.0—12.0 mm. Hab.: Assam . rowlandi

Pubescence variable, but not as above; no FER defined dark hair band restricted
to the middle of tergite 202400 RAI So ac

Apical portion of sternite 7, Beyond fe two hel deci transverse ridges,
at first parallel-sided, then much expanded and terminating in a pair of divergent
flaps separated by a shallow median emargination (Fig. 30, 36). Hind basitarsus
comparatively broad and thin, 3.5 to almost 3.7 times as long as broad, parallel-
sided and strongly laterally compressed; both dorsal and ventral ridges subacute
and hairy, the erect basal hairs of the dense fringe at upper ridge and apex longest,
exceeding greatest width of basitarsus; outer face not sulcate, smooth and shining,
very finely superficially and sparsely punctate, with few short decumbent hairs;
inner face divided lengthwise into a thickly pubescent upper part and a narrower
strip bordering the ventral ridge, which is clothed sparsely with much shorter hairs,
this almost bare lower part being placed at an oblique angle to the dorsal crest (Fig.
27, 33). Supraclypeal area with thick, transverse, yellow bar along epistomal suture;
clypeus and paraclypeal areas almost wholly yellow. Labrum light brown, little
broader than long, anterior border not emarginate, but disk with small midapical
depression, bounded by a tiny crescentic or /\-shaped ridge carrying a small tubercle
on either side; surface rugosely punctate, with tuft of longish hairs not quite
concealing the surface; also a tuft in the antemedian depression. Mandibles dirty
yellow, base and apex obscured. Malar space distinct, not broader than pedicel of
antenna. Ocelli placed in equilateral triangle. IOD : OOD = 77 : 100. Antennal
segment 3 distinctly shorter than 4 + 5; ratio of length to width 100 : 51.5; length
ratios of 3, 4 and 5 = 100 : 75—80 : 75—80. Colour of body vestiture very
variable. Postgradular areas of sternites 4—5 smooth, shallowly impressed, lateral
areas of 4 clothed with long hairs but apical fringe at middle very short; free border
of 5 broadly emarginate, surface with pair of conspicuous diverging patches of
minute, extremely condensed, golden brown decumbent hairs; apex of 6 little
prominent, fringed all round with short radiating hairs (Fig. 29, 35). Body length
13.0—14.0 mm. Hab.: Taiwan and SE China . . . . tainanicola

Apical portion of sternite 7, on either side of the en terne ridges (if
present), at first very broad with strongly projecting side-angles, but soon much
narrowed and tapering towards a variously shaped apex. Hind basitarsus varying in
length, the dorsal ridge sharp, but lower portion gradually swollen from above
down and ventral surface rather convex; outer face whether or not sulcate, inner

8!.

M. A. LIEFTINCK: Review of Asiatic Habropoda 167

face normal, undivided, covered densely all over with uniform setiferous punctures
or minute warts. Supraclypeal area dark brown or black, unmarked. Sternites 2—5
straight-bordered, texture and vestiture of 4—5 not as above . . . . . . 7

Hind basitarsus comparatively narrow in profile, from almost 4 to 4.3 times as
long as broad; its ventral surface either covered all over with bristle-like hairs, or
the naked part is restricted to a linear strip placed at right angles to the opposite
dorsal crest, tapering away and disappearing at some distance from apex. Anterior
face of antennal scape conspicuously yellow... 7 ‘ ts

Hind basitarsus varying in length; its ventral surface, ea the panded side-
edges, with a completely bare, convex central area placed at right angles to the
opposite dorsal ridge, this hairless area variable in width, but extending from base
fopapexs ii. O

Outer face of ind Pri hast or not ere, but vete nen without
any indication of a slightly broadened, convex and naked area. Clypeus, paraclypeal
area and mandibles almost wholly bright yellow. Antennal segment 3 little shorter
thanidad: St... .0, Rea ada ne RO

Outer face of hind ta alloy prg rai vale and somewhat
shining, covered rather sparsely with longish decumbent hairs; ventral surface of its
swollen lower portion slightly convex, bare, and covered ich setiferous warts
towards apex, the impunctate area being reduced to a narrow strip that tapers away
and disappears slightly beyond halfway its length from base (Fig. 46). Mandible
bases, paraclypeal areas, and a thick more or less T- (mushroom-) shaped median
spot on clypeus, bright yellow. Antennal segment 3 a little longer than 4 + 5.
Labrum dark reddish brown, base including the large tubercles lighter brown;
slightly broader than long, anterior border with minute crescentic emargination and
a pair of low, converging anteapical ridges running parallel to the apical border,
each ending in a small tubercle; disk slightly concave, irregularly rugosely punctate,
with long tuft of pale hairs, and a smaller tuft also in the depth of the emargi-
nation. Body pubescence throughout orange-buff, except some dark-tipped hairs
sparsely intermixed above, mostly palest yellow or white on ventral surfaces. Legs
slender; hair fringing dorsal ridge of hind basitarsus much shorter than greatest
width of the latter. IOD : OOD = 53 : 100. Ratio of length to width of antennal
segment 3 = 100 : 40; length ratios of 3, 4 and 5 = 100 : 42.2 : 48.8. Tergite
7 truncated, margin feebly emarginate. Posterior sternal hair fringes pure white, of
great length, dense and partly wavy, directed inward and with recurved apices,
marginal hairs of 4—5 of almost equal length, those of 5 only little shorter
medially than at sides; 6 bluntly triangular, clothed apically with short, stiff
radiating hairs. Sternites 7—8 and genitalia, Fig. 47—48. Body length 12.0 mm
approx. Hab.: Taiwan. . . . + + . bucconis

Legs brown, thin and snito ie and sh dici Saga with relatively
short and pale hairs. Hind basitarsus narrow, parallel-sided, outer face hardly
noticeably concave, not definitely sulcate, surface dull, finely tessellate, clothed with
short, evenly distributed depressed hairs, those fringing dorsal ridge also short
(Fig. 38). Labrum black, somewhat broader than long (100 : 76), widest basally;
disk bounded antero-laterally by a pair of swollen, smooth and shining ridges, close
to the almost straight anterior margin, each of these curving mesiad and ending in a
small tubercle placed on either side of a mid-apical depression; surface flat, coarsely

168

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

rugosely punctate, well visible under rather short, semi-erect pale hairs; apex also
with small tuft in the depression. Malar area distinct, though narrower than length
of pedicel of antenna. Face relatively narrow, less than twice as broad as greatest
transverse diameter of eye at level of supraclypeal suture (100 : 55). Clypeus with
thick, deep black stripe along paraclypeal suture, separating the yellow areas on each
side of it; further down this stripe curves outward, meeting the eye-margin, so as to
fill out the lower edge of the paraclypeal area. Ratio of length to width of antennal
segment 3 = 100 : 44.4; length:ratios of 3, 4 and 5 = 100 : 51.4 : 53.0. IOD :
OOD = 54 : 100. Apical light hair bands of tergites 1—5 narrower, more
contrasting and condensed than in other pale-coloured species, leaving more of the
dark basal portions which are only sparsely clothed with a mixture of long, raised,
brown and yellowish hairs, not at all concealing the somewhat shining integument;
the latter, as in swtepensis, sparsely punctate and finely reticulated. Apex of tergite
7 shallowly emarginate with rounded side-edges (Fig. 39); upper surface clothed
with dense orangish pubescence. Posterior sternal hair fringes relatively short, palest
yellow; sternite 5 shallowly emarginate, apical portion, between the lateral fringes,
with transverse brushy patch of minute, slightly recurved ferruginous hairs; 6 as
described for bucconis. Sternites 7—8 and genitalia, Fig. 40—41. Body length 11.2
mm approx. Female unknown. Hab.: Nepal . . . . . . . apostasia
Legs darker brown and not so slender, tibiae and tarsi Ban long, partly dark
hairs: outer face of hind basitarsus shining, shallowly sulcate, sparsely punctate,
clothed with short depressed hairs; dorsal ridge with compact, very long, black
fringe; expanded ventral ridge strewn with strong, bristle-bearing warts. Hind tibia
brown-haired, but inner and outer faces apically also with long black tufts (Fig. 42).
Labrum reddish brown, growing darker apically, shaped as in apostasia, disk not
ridged laterally, but anterior border with small, almost circular, mid-apical depres-
sion bounded by a low ridge, which on either side ends in a minute tubercle; surface
flat, rugose, rather shining, clothed with short pale pubescence not concealing the
ground; a small tuft also in the mid-apical depression. Malar space parallel-sided,
distinctly broader than length of pedicel of antenna, surface shining, chestnut-
coloured. Face much broader, more than twice as broad as greatest transverse
diameter of eye at level of supraclypeal suture (100 : 44). Clypeus and paraclypeal
area unmarked, save for a small brownish spot (or streak) directed obliquely inward
from the obscured tentorial pit; no dark line bordering paraclypeal suture. Ratio of
length to width of antennal segment 3 = 100 : 42; length ratios of 3, 4 and 5 =
100 : 58 : 60.5. IOD : OOD = 44 : 100. Body pubescence longer and denser,
throughout orange-buff to orange, with brown-tipped hairs sparsely intermixed
upon vertex and occipital area, the abdominal banding broader and indistinctly
limited towards base of tergites. Apex of tergite 7 long and narrow, tip bituber-
culate (Fig. 43), upper surface clothed with long, brownish-black hairs. Posterior
sternal hair fringes palest yellow, considerably longer and denser than in apostasia,
especially the lateral tufts of 3—6, which are very conspicuous, the vestiture of 5
and 6 very nearly like bucconzs, but hair covering 5 more compact at middle and
also distinctly shorter than at sides. Sternites 7—8 and genitalia, Fig. 44—45. Small
species, body length 10.0 mm approx. Hab.: Thailand. . . . . . sutepensis

10.

10%

IR.

12%

M. A. LIEFTINCK: Review of Asiatic Habropoda 169

Hind basitarsus strong, less than three times as long as its breadth at apex, at
least slightly broadened towards apex in profile; hairless area on ventral surface
lanceolate (pelmata), or much broader, definitely sole-shaped. Posterior sternal hair
fringes pale yellow, very long and dense, especially at sides, much as described for
bucconis, but hairs covering sternite 5 much shorter and also more compact than
those of 4, most noticeably so in the middle along on Anterior face of antennal
scape conspicuously yellow . . . DIRE AT

Hind basitarsus slenderer, at least Bice Eire: as an as its gui breadth near
apex in profile and more nearly parallel-sided; hairless area on ventral surface
lanceolate. Posterior sternal hair fringes Lia or white, as before, but hairs
covering sternite 5 not much shorter and more condensed than those of 4 . . 13

Hind basitarsus in profile slightly broadened towards apex, its lanceolate
(torpedo-shaped) naked ventral area light brown, the slightly swollen apex of this
not projecting below and evenly rounded in side view. Legs brown, mainly palely
pubescent, except small apical tufts of dark hair at dorsal ridges of mid and hind
tibiae, and suberect deep black fringe along whole dorsal ridge of hind basitarsus;
dense decumbent pubescence covering inner faces of hind basitarsi pale golden
brown. Hind tibia rather slender, widest beyond halfway length and gradually
expanded towards apex; inner face lacking apical tuft of long hairs (Fig. 68).
Mandible bases, paraclypeal area, antennal scape anteriorly, and a very thick, mush-
room-shaped spot, similar to that of bzcconis, upon middle of clypeus, all pale
yellow; supraclypeal area unmarked, or with pair of tiny transverse yellow streaks.
IOD : OOD = 50 : 100. Antennal scape with external fringe of long pale hairs
(rest of both antennae missing). Apex of tergite 7 truncated (Fig. 69). Sternites
7—8 and genitalia, Fig. 70—71. Size small, body length 11.0 mm approx. Female
unknown. Hab.: Nepal . . . . . . . pelmata

Hind basitarsus in profile distinctly red da en which is produced
below into a bluntly rounded process (Fig. 56); naked ventral area yellow-brown,
conspicuous and well defined, surface undulated, forming a convex sole-shaped
plate (Fig. 57). Legs robust, femora and tibiae rather swollen, clothed with long,
mostly pale, pubescence. Hind tibia more expanded, its greatest breadth at about
midway length, inner face with conspicuous tuft of very long, wavy dark hairs.
Anterior face of antennal scape conspicuously yellow, clothed with longish pale
pubescence anteriorly; segment 3 shorter than 4 + 5. . . . ur 12

Clypeus predominantly black: yellow median spot strongly ei at middle,
rather I-shaped, the transverse anterior bar tapering at either end. Labrum dark
brown, blackened apically, anterior border with distinct crescentic median emargina-
tion. Legs, including basitarsi, dark brown, tarsal distalia lighter; inner faces of
hind tibia and basitarsus, including dense fringe at dorsal crest, black, the sole
almost four times as long as broad, chestnut-coloured, strongly swollen apically.
Dorsal pubescence of thorax and abdomen orange-buff to deep chrome, with dark
hairs sparsely intermixed on thorax, the tergal “banding” fairly distinct, Pioen
sively broader and more distinctly orange posteriorly. IOD : OOD = 52.5 : 100.
Ratio of length to width of antennal segment 3 — 100 : 46.5; length ratios of 3, 4
and 5 = 100 : 58.1 : 62.8. Apex of tergite 7 subtruncated, the sides usually
obtuse-angulate (Fig. 58). Sternites 7—8 and genitalia, Fig. 59—61. Body length
13.8—14.0 mm. Hab.: E Himalaya; Nepal; Assam . . . . . radoszkowskii

170

121%

13:

13%

14.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Clypeus predominantly yellow: median spot thick, less constricted, rather diabolo-
shaped; dark lateral marks brownish and not sharply outlined mesially. Labrum
light brown, anterior border slightly crenulated at middle, not emarginate. Legs,
including basitarsi, orange-buff, the femora and tibiae slightly darker; pubescence
throughout warm orange, except the long inferior apical tuft at hind tibia, which is
black, and a thin fringe at dorsal crest of hind basitarsus, brown; sole a little over
four times as long as broad, light brown, slightly less expanded and swollen apically
than in radoszkowskii. Dorsal pubescence of thorax and abdomen longer, more
condensed and more brightly orange, the tergal “bands’’ deepest in tint posteriorly.
IOD : OOD = 60 : 100. Ratio of length to width of antennal segment 3 =
100 : 51; length ratios of 3, 4 and 5 = 100 : 65.7 : 71.4. Apex of tergite 7
narrower (Fig. 64, unique male). Sternites 7—8 and genitalia, Fig. 65—67. Body
length 12.0 mm approx. Hab.: Assam . . . 2) 2) plantifera

Clypeus predominantly brownish black, elle made band more or less
I-shaped, strongly constricted and considerably narrower than dark lateral parts,
clothed abundantly with very long, suberect dark bristles and decumbent pale
converging hairs. Legs dark brown; pubescence pale, except conspicuous fringes
alongside lower surface of hind tibia and a dorsal tuft at apex of same, these hairs
long, raised, and predominantly or wholly black. Hind basitarsus brownish black,
not at all broadened towards apex in profile; outer face shining, longitudinally
sulcate, sparsely hairy; hairless ventral area dark brown, strongly convex, surface
dullish, with few superficial punctures; a dense hair fringe along full length of
dorsal ridge, jet black. IOD : OOD = 57.5 : 100. Anterior face of antennal scape
conspicuously yellow, with external fringe of long pale hairs; 3 a little shorter than
4 + 5, ratio of length to width 100 : 38; length ratios of 3, 4 and 5 = 100:
50.5 : 60. Tergite 7 somewhat attenuated, but apex truncated (Fig. 51). Sternites
7—8 and genitalia, Fig. 52—53. ad length variable, 13.0—14.5 mm. Hab.: SE
Chinas (Eukien) nn 2% . . . imitatrix

Clypeus predominantly dir an cda ila Eos doo widest below, but
slightly narrower than greatest width of paraclypeal area, at either side of clypeus,
and a pair of more or less isolated brown discal spots, placed a little more inward,
slightly beyond halfway length from base. Anterior face of antennal scape entirely
dark brown, unmarked with yellow; 3 subequal to, or a little longer than, 4 + 5.
Legs light to dark brown, pubescence not quite as described above, mainly pale;
raised hair covering lower surface of hind tibia short, sparsely distributed and pale-
coloured. Hind basitarsus more slender, subparallel-sided; outer face sulcate, rather
shining, sparsely pubescent and punctate; hairless area narrowly lanceolate, light
brown, otherwise as in pelmata (Fig. 72, 78, 85). = er palely pubescent,
lacking black hairs. . . PR al 4

Head large and face ral Bread: dine Fe inner orbits and
diameter of eye at level of supraclypeal suture in the ratio of 100 : 50. Protruding
median portion of clypeus broadest well beyond halfway its length from base;
length : breadth ratio of clypeus in frontal view about 90 : 100. Clypeus with
yellow lateral streak bordering anterior end of paraclypeal suture. IOD : OOD =
51 : 100. Ratio of length to width of antennal segment 3 = 100 : 40; length ratios
of 3, 4 and 5 = 100 : 46.5 : 51.2. Second submarginal cell markedly longer than
high, receiving recurrent nervure well before its apex. Apex of tergite 7 narrow,

141.

M. A. LIEFTINCK: Review of Asiatic Habropoda 171

hind margin very shallowly concave. Sternites 7—8 and genitalia, Fig. 73—76.
Size a little larger and stature more robust. i length 13.0—13.5 mm. Hab.:
West Himalaya . . . ee. deiopea
Head comparatively cal) nd face narrower: dee quia inner orbits and
diameter of eye at level of supraclypeal suture in the ratio of 100 : 56—57. Pro-
truding median portion of clypeus broadest at or a little beyond halfway its length
from base; length : breadth ratio of clypeus in frontal view about 110 : 100. Yellow
lateral streak on clypeus bordering anterior end of paraclypeal suture present
(Nepal), or wanting (Darjeeling). IOD : OOD .= 60 : 100. Ratio of length to
width of antennal segment 3 = 100 : 40—42; length ratios of 3, 4 and 5 =
100 : 42 : 47.4 (Nepal), or 100 : 44.4 : 55.5 (Darjeeling). Second submarginal
cell almost square, recurrent nervure entering cell a little before its apex. Apex of
tergite 7 narrow, truncated (Fig. 79, Darjeeling), or as in Fig. 86 (Nepal). Sternites
7—8 and genitalia, Fig. 80—83 (Darjeeling) and Fig. 87—89 (Nepal). Averages
smaller in size, stature more slender. Body length 11.0—12.5 mm. Hab.: East
AA an SP Nepal NRR es apatelia

KEY TO THE SPECIES, FEMALES
N.B. — The females of H. apostasta, pelmata and plantifera are still unknown

At least tergites (1) 2—4 with complete, well defined, pale (sub)apical pubes-
cent bands or lines, which are about equally broad, consisting of closely-set decum-
bent hairs entirely concealing the marginal surface. Outer hind tibial and/or
basitarsal scopal hairs never quite black . . . . HAE

Tergite (1) 2—4 diffusely pubescent, Benel vanieoloured: Hae lacking
contrasting, clearly defined, apical hair bands of one colour; hind margins of
integument often broadly pale, membranous and seemingly “banded” . . . 5

Eyes not enlarged, converging below, only little shorter than upper interorbital
distance (100 : 106); inner orbits nearly straight, vertex at summit a little broader
than lower interorbital distance. Ocelli placed in a broad, isosceles triangle; distance
separating lateral ocelli two or more times their own diameter; interocellar (IOD)
and ocellorbital (OOD) distances equal (hence IOD : OOD = 100 : 100). Mouth
parts, face and antennae all black, except basal tubercles of labrum and mid portion
of mandibles diffusely brownish. Labrum slightly broader than long, disk with
conspicuous dense tuft of decumbent pale hairs. Malar space shorter than pedicel of
antenna; antennal scape lacking anterior fringe of long hairs; segment 3 subequal
to 4 + 5. Front of head moderately protuberant. Clypeus rather convex, protruding
anteriorly beyond eye in profile by as much as width of latter; surface rather closely,
irregularly punctate and hairy, especially at sides; epistomal suture in frontal view
straight. Inner ramus of mid and hind tarsal claws almost half as long as outer;
arolia well developed. Abdominal tergites closely reticulate-punctate; light apical
bands fairly broad, occupying at least one-fourth of the exposed surface; much
longer raised pale hairs present on rest of tergite 1, but very short, sparsely
distributed, more or less appressed and mostly black, on the exposed basal portions
of remaining tergites. Sternites 2—5 with dense subapical fringes of suberect pale
hairs, not much longer laterally than at middle. Pygidial plate broadly tongue-
shaped, much as in rowlandi (Fig. 26), but even broader, more gradually tapered,

172

20:

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

almost flat, lacking depressed margins; surface finely transversely striated, the striae
convex posteriorly. o species, body n 15—16 mm approx. Hab.: W and
NE China Gon: . + +. pekinensis

Eyes larger and more le, Si alle or ed so, inner ANG straight, vertex
at summit subequal in width to lower interorbital distance. Ocelli placed in an
almost equilateral triangle; IOD much shorter than OOD. Labrum dark, fully twice
broader than long, widest basally, surface rugose; pale hairs on disk shorter and
less dense, not concealing surface. Malar space broadest anteriorly, about as long as
pedicel of antenna. Front of head broad, ‘‘pug-nosed’’, conspicuously bulging, but
clypeus evenly convex, protruding anteriorly beyond eye in profile by at most 34
(usually less) the width of eye, its surface smooth, shining, almost bare, strongly
punctate and with conspicuous yellow marks (Fig. 8—10); epistomal suture
distinctly arched in frontal view. Antennal scape lacking anterior fringe of long
hairs, unmarked with yellow. Claws slender, inner ramus of mid and hind pairs at
least half as long as outer; arolia well developed, of large size. All abdominal
segments closely punctate; light apical tergal hair bands variable; pile on disk of
tergite 1 raised and considerably longer than on following segments, though not
concealing the surface. Pale suberect hairs fringing apical margins of sternites 2—4
not noticeably long and wavy, though longest at sides of 3—4. Pygidial plate
brown, very broad at extreme base (exceeding total length), then tapering rapidly
to a narrow and almost pointed tip, the sides distinctly incurved, margins a little
upturned; surface slightly convex, rather shining, a tessellated. Smaller-sized
Species un. 3

Apical tergal kart: bands te: cad ad Dae cengia Abe one- ri
or more of the exposed surface of each tergite, colour pale orange-yellow, hairs
curled and almost as crowded as in male; apical band of tergite 5 cinnamon brown,
projecting broadly and triangularly basad at middle, the long hairs at sides pale;
integument of tergites almost black, surface dull, faintly lustrous, evenly, extremely
closely and rather deeply punctate, diameter of punctures exceeding interspaces;
punctation equally dense on pregradular areas of sternites, but more sparse and
superficial on postgradular portions, except along hind margins. Outer hind tibial
scopal hairs pale orange-yellow, not quite concealing surface, outer faces of hind
basitarsi much more sparsely hairy and darker; penicillus brown. Anterior border
of labrum but little convex, median emargination very small; disk lacking dense
hair tuft, but apex with longish, pale golden fringe. Clypeus chrome, marked with
pair of large, diverging, subrectangular brown patches, closely approximated or
even fused together at extreme base and extending to about 24 to 34 length from
base; these marks are obliquely cut off anteriorly, longest at each side of the median
line, so as to save a tripartite yellow mark bordering the clypeus anteriorly (Fig. 8).
Paraclypeal area black, clothed densely with appressed yellowish white hairs.
Supraclypeal area frequently also with trace of yellow bordering epistomal suture.
IOD : OOD = 37.5 : 100; distance separating lateral ocelli subequal to their own
diameter. Eyes longer than upper interorbital distance (100 : 78). Antennal segment
3 almost twice as long as thick and equal in length to 4 + 5 (length ratios of 3, 4
and 5 = 100 : 45.5 : 54.5). Marginal cell of fore wing longer than distance from
its apex to wing tip. Pygidial plate (Fig. 11) reddish brown, abruptly changing to
blackish brown at the constriction, basal portion strongly convex, surface shining,

3E.

41,

M. A. LIEFTINCK: Review of Asiatic Habropoda 173

finely transversely reticulate-striate, except apically. Body En variable, 12.3—
14.0 mm. Hab.: E China . . . . . . sinensis
Apical tergal hair bands linear ned Ess Ean oen fateh less than one-
fifth of the exposed surface of each tergite, colour pale chrome; integument of
tergites glossy black, almost bare, evenly finely punctate, diameter of punctures
smaller than interspaces. Outer hind tibial scopal hairs bright orange-rufous, entirely
concealing surface; outer faces of hind basitarsi, including penicillus, mainly dark-
haired, the inner ones black. Marginal cell of fore wing only slightly longer than
distance from its apex to wing tip (100 : 91.6—92.3). Males unknown .
Supraclypeal area wholly brownish black; clypeus with inverted T-shaped, chrome-
yellow mark, Fig. 9, whole surface shining, distinctly uneven (rather rugose),
irregularly and scatteredly punctate, except basally, where punctures are more
crowded and wider than the interspaces; supraclypeal area more densely and deeply
punctate, most punctures confluent at sides, those on summit of head smaller and
more widely spaced. Pale posterior bands of tergites 1—4 linear, about 0.2 mm
broad, composed of finely plumose hairs. Outer hind tibial scopal hairs throughout
orange-rufous, those of basitarsus relatively long, dark brown only along dorsal
ridge, and of penicillus, the discal hairs mainly light orangish. Pygidial plate, Fig.
12. Body ui 13.0 mm approx. Hab.: Indochina (South Viet Nam)
: tumidifrons
eye area ik Procal Sie eel vallo crescent bordering the
suture; clypeus with conspicuous anchor-shaped chrome-yellow mark (Fig. 10);
whole surface smooth and shining, but with fewer punctures than in tumidifrons,
even at base smaller and more widely spaced. Pale posterior bands of tergites 1—4
all a little broader, about 0.3 mm or more, otherwise as described above. Outer hind
tibial scopal hairs generally shorter, mainly orange-rufous, but becoming more
brownish towards ventral ridge, those covering basitarsus distinctly shorter and more
numerous than in zumidifons, very dark brown, almost black. Pygidial plate, Fig. 13.
Body length 13.0 mm approx. Hab.: Thailand . . . . . . . . orbifrons
Body pubescence dense, varicoloured: alternately yellow, black and orange, these
coloured areas transverse, broad and sharply delimited; if tergites are predominantly
black-haired, then pubescence on dorsum of thorax also mainly black. Integument of
abdominal tergites not pallid and membranous apically. Outer faces of femora and
tibiae black-haired; tarsal arolia vestigial. Labrum much broader than long. Disk of
clypeus only slightly convex, closely punctate, almost bare. Front of head mainly
black and dark-haired. Ocelli placed in equilateral Cae distance separating
lateral ocelli less than twice their own diameter. . . à A ee NG
Body pubescence usually less dense and not colonel as done: if tergites are
varicoloured, then the coloured areas are not sharply delimited. Hind margins of
abdominal tergites mostly broadly pallid. At least outer faces of hind tibiae and/or
basitarsi light-haired; tarsal arolia well developed. . . . . MEN ES
Eyes large and Fute longer than upper interorbital distinte, inner orbits
subparallel, but distinctly convex in frontal view. Clypeus much less protruding
anteriorly beyond eyes in profile than half diameter of eye (100 : 37). Labrum
subrectangular, ratio of length to width = 60 : 100, not projecting anterad, border
almost straight but considerably thickened medially, with deep U-shaped notch and
tuft of ferruginous bristles in the depth of the emargination, apex not preceded by

174

Tip:

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

a subparallel ridge. Dorsum and sides of thorax clothed evenly with rather short,
pale yellow to orange pubescence; mesonotum with a thick, transverse, black median
band or patch between the wings. Abdominal pubescence also comparatively short,
on tergite 2 (save for a narrow light apical fringe) black, on 1 orange-yellow and
black, on succeeding tergites variable. Pygidial plate (Fig. 14—15) broadly
triangular, tapering gradually towards apex, which is bluntly rounded; sides slightly
concave, margins a little thickened and raised; surface dull, markedly convex, finely
transversely striate; colour reddish-brown, apex obscured. Robust, short-legged,
compactly built species . . SUSA

Eyes much smaller and less peominent, si. ion onee: nen upper inter-
orbital distance (100 : 88), inner orbits subparallel and almost straight in frontal
view. Clypeus protruding anteriorly beyond eyes in profile for almost half diameter
of eye (100 : 46). Labrum broad, ratio of length to width = 68 : 100, projecting
more markedly anterad than in male, apex obtuse-angulate, anterior border only
slightly thickened medially, but preceded by a low arched ridge, narrowly inter-
rupted in the median line, which runs subparallel with free margin. JOD : OOD =
70 : 100. Antennae, including scape anteriorly, dark brown; segment 3 much longer
than 4 + 5, length ratios of 3, 4 and 5 = 100 : 30 : 40. Head dark brownish black
or black, except that basal tubercles of labrum are light brown and mid portion of
mandibles chestnut. Legs blackish brown, black-haired, the tarsi lighter brown; hair
on outer faces of basitarsi reddish brown to black, on the inner golden brown to
ferruginous; dorsal fringe of mid and hind basitarsi black; penicillus brassy. Dorsal
pubescence longer than in the two preceding species, pattern much as in male,
except that the long dense canary yellow hairs (raised on 1, decumbent on 2)
occupy almost whole surface of 2 and are tipped with black; 3 entirely black, 4—5
orange-rufous, the fringe at apex of 5 very brightly so. Sides and ventral surfaces
mainly dark brown, except long white-tipped hairs at base of fore femora, and long
dense tufts covering most of the genal area and thoracic pleurae below wings, which
are silvery white. Pygidial plate (Fig. 21) broadly triangular, tapering towards
apex, which is bluntly rounded; sides almost straight, margins a little upturned;
surface dull, evenly and but slightly convex; colour reddish-brown, growing darker
apically. Body length 15.5—16.0 mm. Hab.: Assam . . . . . turneri

Black pubescent band between fore wings about as broad as sn yellow mesonotal
area in front of it and only little broader than the yellow patch covering posterior
portion of thoracic dorsum; colour of light thoracic pubescence bright yellowish
tawny, on tergite 1 slightly more orangish, except at posterior margin of the latter,
where it is occasionally black; 2 totally black-haired, save for a well-defined apical
fringe of orange; 3—4 wholly bright orange, the hairs on basal portion raised, those
along posterior margins longer, dense and decumbent; 5 dark-haired at base, the
broad apical fringe of long dense hairs ferruginous. Long semi-erect hairs fringing
hind margins of sternites 2—4 light yellow, those at 5 pale ferruginous. Black
plumose hind tibial scopal hairs relatively short, not exceeding greatest breadth of
tibia. Front of head brownish black or black, supraclypeal area with crescentic or
triangular yellow spot bordering frontoclypeal suture. Pygidial plate, Fig. 15. Body
length 15.5—16.5 mm. Hab.: SE China (Fukien) TAPER Se . . . mimetica

Black pubescent band between fore wings about twice as Toni as the orange
mesothoracic collar in front of it and at least 11/, times as broad as the orange band

KO?

M. A. LIEFTINCK: Review of Asiatic Habropoda 175

covering posterior portion of thoracic dorsum; scutellum and upper part of sides
below wings brightest orange, the pubescence lower down acquiring gradually a
yellowish tint. Only basal half of tergite 1 orange, for the rest black-haired; 2—4
entirely black, except narrow fringes of orange at sides of 1 and along full length
at posterior margins of 2—4, the consistence of pilosity as in mimetica; 5 black-
haired at base, the broad apical fringe of long dense hair bright golden-brown.
Long raised hairs fringing hind margin of sternites all black. Black plumose hind
tibial scopal hairs conspicuous, much denser, and longer than greatest breadth of
tibia. Front of head as in mimetica, but supraclypeal area with well defined creamy-
yellow crescent bordering the suture. Pygidial Po Fig. 14. Body length 15.5 mm
Hab.: Indochina (North Viet Nam). . . . . . disconota
Vestiture of thorax, abdomen and genal area buff yellow see much as in
turneri; hair covering mouthparts, face, antennae and summit of head mainly dark
brown. Dorsum of abdominal tergite 2 with transverse, ill-defined, but conspicuously
blackish brown band of light-tipped hairs; dense appressed pubescence covering
tergite 5 and sides of 6 ferruginous to ochraceous-orange; outer faces of mid and
hind tibiae and tarsi, including penicillus, similarly coloured. Head, including
labrum, clypeus and antennae, throughout brownish-black to black, otherwise much
as in male, mandibles dark chestnut at middle; clypeal disk somewhat shining,
covered densely with partly coalescent punctures. Labrum longer than in twrnert,
anterior border scarcely ridged before margin, with minute crescentic median
emargination. Clypeus protruding anteriorly beyond eyes in profile for more than
half diameter of eye (100 : 80); surface rather shining, closely striato-punctate,
there being few isolated punctures. Ocelli placed in isosceles triangle, distance separ-
ating lateral ocelli exactly twice their own diameter; IOD : OOD = 75.6 : 100.
Antennal segment 3 only little longer than 4 + 5, length ratios of 3, 4 and 5 =
100 : 40.5 : 45.2. Pygidial plate (Fig. 26) chestnut-colour, very broad, the
converging sides almost straight; surface smooth, disk very slightly convex,
microscopically tessellate, margins broad, thin, and a little as apex bluntly
rounded. Body length 13.0—13.5 mm. Hab.: Assam. . . . . . rowlandi
Vestiture of thorax and abdomen very variable, but Bel is no poorly defined
dark hair-band restricted to the middle of tergite 2. Clypeus and supraclypeal area
more strongly protruding from face than in disconota, mimetica, rowlandi and
turneri, arising abruptly in a short step from the deeply recessed antennal sockets
and flat upper portions of the paraclypeal areas. Pygidial plate not as above . . 9
“Similar to male, but larger and more robust. Dense hair of abdomen very red.
Clypeus convex, the upper third in middle with a shining keel, from the lower end
of which starts a creamy-white vertical band, rapidly broadening below to form a
broadly triangular pale area; labrum reddish; mandibles with more than the basal
half pale rufous; malar space well developed; outer side of hind tibiae with rich
copper-red hairs; hind basitarsi with black hair. The cheeks below have long pure
white hair.” (Taken from i description). Other features not known. Hab.:
railand sie op: . . sutepensis
Combined characters do as bone: Ale Leni or a: feeble keeled basally.
Dense hair covering abdomen, if at all partly reddish, then only so on posterior
tergites. Outer faces of hind basitarsus with few dark hairs interspersed . . 10
Pygidial plate (Fig. 32) elongate, tongue-shaped, distinctly longer than its

11.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

greatest width at base, downcurved and gradually narrowed towards the end; sides
almost straight and apex simply rounded; surface even, not at all keeled or ridged,
finely, superficially and transversely striated from base to apex; colour black, turning
to red-brown towards base. Vestiture of thorax and abdomen nearly always partly
obscured (or even black) dorsally, but apical segment(s) of abdomen invariably
orangish or rufous. Outer faces of hind tibia and basitarsus clothed with long, bright
orange-chrome to vivid orange-rufous scopal hairs. Integument of head above and
clypeal areas unicoloured brownish black or black, clypeus occasionally with ill-
defined rusty brown median dot at some distance from apex; disk shining and for
the greater part almost bare, the sides only clothed with soft decumbent pubescence
interspersed with long, erect, dark bristles; whole surface evenly, superficially
(though rather closely) punctate, all punctures isolated and smaller than the
microscopically reticulate interspaces; occasionally a narrow, almost polished and
impunctate, median area. Labrum and most of the mandibles red brown to chestnut;
labrum broader than long, much as in male: with pair of closely approximated, sub-
triangular tubercles placed behind anterior border, which itself is scarcely excised
medially. IOD : OOD = 66.6—72 : 100. Antennal segment 3 much longer than
4 + 5. Body length 14.0—15.5 mm. Hab.: Taiwan and SE China . . tainanicola

Pygidial plate shorter, more nearly triangular, only little longer than its greatest
width at base, where it is very broad, but then tapers rapidly, the sides frequently
curving inward and converging to a narrow and pointed tip. Vestiture of thorax
and abdomen of uniform colour, never obscured, varying from pale ochraceous to
deep chrome, the long erect or suberect dorsal hairs often tipped with brown. Scopal
hairs on outer face of hind tibia and basitarsus less brightly coloured (occasionally
silvery yellow). Clypeus usually more dullish, whole surface densely, rugosely
striato-punctate, there being no isolated punctures; colour variable, but nearly always
with yellow median spot before the apex. Labrum shaped much as in fainanicola,
but anteapical tubercles larger, less approximated and forming together a wide V-
shaped arc, hence upper anterior border distinctly emarginate; mandibles for the
greater part pale yellowish. Antennal segment 3 longer than 4 + 5 . . . . 11

Clypeus and supraclypeal area dull, uniformly, rather closely, rugosely punctate,
all punctures confluent; whole clypeal surface clothed evenly with longish, decum-
bent, pale silvery yellow hairs concealing part of the surface; ground colour brown,
except a transverse, ivory yellow band, broadest at middle, just before the apex.
Labrum only little broader than long, brown, becoming almost black apically, the
basal tubercles lighter; subapical teeth black, triangular and flattened, separated
from each other by a crescentic emargination. Mandibles for the greater part ivory
yellow, the distal portion becoming brown. First three antennal segments brown,
rest missing. Vestiture generally pale (discoloured ?), uniform pale ochraceous,
lacking black hairs; legs brown, the external scopal hairs pale silvery-yellow, almost
white. Abdomen blackish-brown, all segments feebly “banded’” posteriorly, these
bands composed of pale appressed pubescence, the disks moreover with abundant,
somewhat darker, erect hairs; dense patch covering tergite 5 pale ochreous, the
fringe of stiff lateral bristles surrounding the pygidial plate of 6 still darker,
ferruginous-brown. Long posterior sternal hair-fringes palest yellow, almost white.
IOD : OOD = 60 : 100; distance separating lateral ocelli slightly less than twice
their own diameter. Pygidial plate (Fig. 55), chestnut-coloured (only partly

112.

12:

12%:

13.

M. A. LIEFTINCK: Review of Asiatic Habropoda dy

exposed), feebly convex, its median portion raised into a distinct, broad, longi-
tudinal ridge, whole surface finely transversely reticulated. Body length 13.0 mm.
Male unknown. Hab.: West Himalaya. . . . MT AC" hookeri

Clypeus and supraclypeal area rather shining, dos er and irregularly
striato-punctate, the elevated and finely tessellated rugae on the basal portion more
closely set than the smooth and broader ridges on the distal area, the latter glossy,
slightly sloping and feebly triangular in outline, with faint indication of an
impunctate median ridge; central area of clypeus bare or sparsely hairy; ground
colour light to dark brown, frequently with yellow median spot, but lacking a
transverse yellow bar just before the apex. Vestiture generally deeper in tint and
brighter, especially the scopal hairs and abdominal segments posteriorly; long erect
or suberect hairs generally darker than the closely appressed pubescence, which
spreads in a diffuse pattern over the surface of all tergites, this tomentum becoming
progressively more dense, compact and vividly coloured (orange-buff to deep
chrome) towards the end of abdomen. me species of small to medium
MAN oe Fan T2

Larger a boa length 13. 0—16. 0 mm. Hind Pis su admixture of
dark hairs at least basally and along upper and lower margins. Pygidial plate (Fig.
49, 54, 62) gently downcurved, reddish, the apex obscured, surface finely trans-
versely striated. Pale tufts of soft, finely branched hair at postero-lateral margins
of sternites 3—5 much longer than the Re fringes, which are also of a
darker yellow tint . . ERN LS

Smaller species, body gli 12. (SIC 0 mm. No dire of Blade hairs on
outer face and margins of hind basitarsus. Pale tufts of soft, finely branched hair
at postero-lateral margins of sternites 3—5 longer than the 2 rana fringes,
but all are of the same light colour . . Neo)

Clypeus and supraclypeal area dark Blackish brt rich a dich tinge, clypeus
shining, with well-defined ivory spot placed upon middle of distal half, just before
anterior border, this spot shaped like an isosceles triangle, the base of which is
shortest and a little convex, the sides concave, the apex acuminate and pointing
basad. Malar space as long as pedicel of antenna. Raised and appressed hairs
covering dorsum of thoracic segments and abdomen moderately long and dense,
neither quite concealing the black ground of thoracic dorsum, nor the disks of
tergites 1—4; dorsal pubescence of thorax sallower (greyish-ochraceous) than that
of abdomen. Dense hair covering tergite 5, as well as the scopal hairs of hind
basitarsal penicillus, golden yellow; upper and lower borders of hind basitarsus with
distinct basal admixture of almost black hairs. Pile upon tergite 5, on either side of
pygidial plate, golden brown; plate with feebly indicated median ridge. IOD : OOD
— 50 : 100; distance separating lateral ocelli less than twice their own diameter.
Length ratios of antennal segments 3, 4 and 5 = 100 : 30 : 40. Pygidial plate
(Fig. 62). Slightly smaller, 14.0—15.0 mm. Hane East Himalaya Nepal; Assam
radoszkowskii

Glyfieus aac edi area dot so dee pen dullish and more closely
striato-punctate, yellow median spot before anterior border variable. Malar area a
little longer than pedicel of antenna. Raised and appressed hairs covering dorsum
of thoracic segments and abdomen longer, uniformly more dense and brightly
coloured, all tergal hairs concealing most of the surface, even of tergites 1 and 2.

178

14.

141.

15:

192:

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Dense hair covering tergite 5, as well as the scopal hairs of hind tibia, more vividly
orange; hair on outer face of hind basitarsus somewhat darker than on tibial scopa,
but not blackish at base; penicillus golden orange. Pilosity of tergite 6, on either
side of pygidial plate, as in radoszkowskit. Length ratios of antennal segments 3, 4
and 5 = 100 : 28—30 : 40—40.3. ane more robust and larger, 14.5—17.0
mm NUM EL
Gippea! Cho: nee ad! Mich as in ii ‘but ies sharply
defined, the apex bluntly pointed. Pygidial plate (Fig. 49), with well pronounced
median ridge. Ocelli larger, more closely approximated, distance separating lateral
ocelli scarcely greater than their own diameter; IOD : OOD = 54.3 : 100. Body
length 15.0 mm approx.sHabawaiwan- . tr teater A cee wey buccoms
Clypeal spot still smaller, diffuse, usually elongate-oval, and occasionally
vestigial, but rarely absent altogether. Pygidial plate (Fig. 54), with median ridge
unapparent. Ocelli smaller, more widely apart, distance separating lateral ocelli less
than twice their own diameter; IOD : OOD = 70 : 100. Size variable, 14.5—16.0
mis ab: SE Chinas een . . imitatrix
Mussoorie: Pygidial plate Ge 77) chestnut oie dark brown apically,
shaped much as in radoszkowskii, the median ridge broad but rather low, and fine
transverse striation effaced (worn ?). Protruding median part of clypeus distinctly
longer than broad in frontal view. Front of head, including labrum and clypeus,
very dark brown, almost black, clypeus without any indication of a yellow mid-
apical spot, surface coarsely, rugosely punctate, basal three-fourths with low, ill-
pronounced, irregular median ridge. Labrum with base of converging ante-apical
ridges impunctate. Mandibles dirty ochreous tipped with black; malar area slightly
longer than pedicel of antenna. Galea of proboscis short, 2.2 mm. IOD : OOD =
64 : 100. Antenna brownish black, the flagellar segments 5—12 distinctly light
brown anteriorly; length ratios of 3, 4 and 5 = 100 : 30 : 37.5. Wing margin
somewhat lacerated; second submarginal cell markedly longer than high, receiving
recurrent nervure shortly before apex. Raised hairs covering dorsal thoracic segments
and first two gastral tergites greyish yellow, many hairs on mesoscutum anteriorly,
and on distal parts of tergites 1—2, more or less obscured. Scopal hairs orange-buff
to orange; penicillus golden brown. Hab.: West Himalaya . . . . . deiopea
Darjeeling: Pygidial plate (Fig. 84) with convex median ridge occupying most of
the surface, apex attenuated and almost pointed. Protruding median part of clypeus
a trifle broader than long, almost square in frontal view. Front of head, including
labrum, proboscis and clypeus yellowish brown (ochraceous-buff), clypeus moreover
with tiny, ill-defined, roundish yellow dot upon middle of distal half before anterior
border, its surface less coarsely punctate; ante-apical ridge and free margin of
labrum black. Mandibles yellow, apices obscured; malar area as before. Galea of
proboscis longer, 2.7 mm. IOD : OOD = 53.5 : 100. First three segments of
antenna dark brown, rest of flagellum ochraceous anteriorly, dark brown posteriorly;
length ratios of 3, 4 and 5 = 100 : 32.2 : 40. Wing margin entire; second sub-
marginal cell almost square, recurrent nervure interstitial, Pubescence more vividly
coloured, distinctly more orangish, similar to fresh males, many hairs dark-tipped,
but abdominal pile brighter; hair at sides of thorax and of body parts underneath
palest yellow, not pure white. Scopal hairs much as in deiopea, but more yellow-
orange with distinct golden shine; aa slightly darker. Hab.: East Himalaya
: u oe apatelia

M. A. LIEFTINCK: Review of Asiatic Habropoda 179

DISCUSSION OF SPECIES AND SUBSPECIES

Habropoda pekinensis Cockerell

Habropoda pekinensis Cockerell, 1911, Proc. U.S. Nat. Mus., 39: 642—643 (4 2 Pekin, China,
1901, M. L. Robb) — Lieftinck, 1966, Tijdschr. v. Ent., 109: 144 (full references; addit. notes).

Material examined. — North China: 1 9, labelled “Tjinen Cho [Tshan
Ghan?} (2) Apr. 13, ’29”, in Cockerell’s hand, and “H. pekinensis, det. T. D. A.
Cockerell, pres. by Imp. Bur. Ent. Brit. Mus. 1929-570” (print) (general coll., BM).

For the synonymy of this bee, see my previous remarks (1966). Since briefly reporting
on pekinensis, I could examine a series of both sexes in the late V. B. Popov’s collection
at Leningrad; these are from eastern USSR provinces and bear locality labels in Russian
characters. Though none of them were identified with alashanica Gussakowskij, the
females agreed with Cockerell’s specimen of that sex in the Brit. Mus. (Nat. Hist.)
collection. One of the males from “Pekin” in the National Museum of Natural History
(USNM) was selected by Cockerell as the type of H. pekinensis.

As the salient male characters of pekinensis can not be described or illustrated at
present, I am merely quoting the original description of both sexes.

Original description. — “Male. Closely related to H. zonatula Smith; on comparison
with a zonatula from Smith’s collection (Nicopolis, May, 1836) the following differences
are found: A little less robust; abdomen narrower, less triangular, more as in Antho-
phora; pubescence paler, not so red; flagellum longer and more slender, fourth antennal
joint much longer; first 7.7. not quite reaching apex of second s.m.; black on clypeus
reduced, the middle broadly yellow to top, the large black markings variable, but con-
stricted in middle; pygidial plate much broader; anterior femora strongly keeled beneath
at base; hind tarsi red. The anterior coxae have the characteristic long backwardly-
directed spines, and the hind basitarsi the great flattened lamina. The male is the
type. — Female. About 16 mm. long, with the same ochreous hair covering thorax,
the same black abdomen with light hair-bands; face, mandibles, and antennae entirely
dark; fifth abdominal segment with the hair clear fox-red in middle, cream-colored at
sides; scopa of hind legs light golden-ferruginous; eyes pale green, stained with red. —
Habitat. Pekin, China, 1901 (M. L. Robb). Four males; April 19 (2), April 20, April
21. Twenty-three females; April 19 (12), April 20 (5), April 21 (5), April 22 (1). —
Type. Cat. no. 13426, U.S.N.M.” — Follows a comparison with a number of unrelated
species.

This robust species clearly belongs to a group of six closely interrelated western
Palaearctic Habropoda ranging from the Mediterranean far into continental Asia. It is
the only member that occurs so far north-eastward. H. pekinensis is treated here only to
demonstrate some differential characters over against all other Asiatic species having a
more south-eastern distribution; it will be seen that this section comprises a much less
homogeneous assemblage of forms. It comes nearest the Mediterranean H. zonatula Smith
and a few other species inhabiting central West Asia, sharing with these the more
distinctive features of Habropoda in its restricted sense. The male is, indeed, closely
similar to zonatula, but I noted a difference in the lobes at the hind basitarsi, which
in pekinensis are considerably larger, broader, and also lighter in colour. The male
internal structures could not be examined. For some further details of the female, see
the descriptive key; the pygidial plate is shaped similarly to that of zonatula.

180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Habropoda sinensis Alfken
Fig. 1—3, 7—8, 11
Habropoda sinensis Alfken, 1937, Ent. & Phytopath., 5 no. 20: 404—405 (9 & Chekiang, Tien-
Mu-Shan, 24.viii.1936, T. C. Maa) — Woo, 1965, Chinese Ins. Econ. Importance (83 pp., 7 col.

pls.), 9 Hym. Apoidea: 66 (6 2 Chekiang, Kiangsi and Szechuan) — Lieftinck, 1966, Tijdschr. v.
Ent., 109: 147 (note).

Type material examined. — China: 1 9 (holotype, no. 204), 1 4 (allotype),
West Chekiang, Tien-Mu-Shan, 24.8.36, Tsing-chao Maa; 1 G 1 9 (paratypes), with
same locality information and all bearing red Typus-, Allotypus- and Paratypus-labels,
in J. D. Alfken’s handwriting (MNB).

Further material examined. — China: 1 & 1 9, Kuling, Musée Heude, 7.7.35,
O. Piel, identified with ‘“? Habropoda sinensis Alf. 9 g'” (unknown hand), under
drawer-label of V. B. Popov: “Emphoropsis sinensis Alf.” (ZIL). Long series of both
sexes (33 g', 51 9) from various localities in Fukien Prov., as follows: Shaowu distr.:
Tachulan, 1000 m and Wuku, 350 m; Chungan distr.: Bohea Hills, 600 m; Kuatun,
1600-2000 m; Sanchiang, 600 m; Kienyang distr.: Hwangkeng, 500 m; various dates,
from vi-x.1942-1945, all T. C. Maa (ML and various other collections).

With the two new species, H. tumidifrons and orbifrons, defined hereafter and in the
descriptive key, sinensis forms a fairly distinctive section of the genus. The key characters
will suffice to recognize them from all other Habropoda; for additional information
on colour and pubescence, see also the original description. The bees of this little group
are mainly distinctive by their more compact build, short antennae, bare and shiny
abdomen, and also by the narrowly banded tergites and much shorter pubescence, re-
calling Amegilla in general appearance. The group is characterized further by the large
head, broad, “pug-nosed” face and striking body punctation. These features, shared
alike by all three, point to close interrelationship. As to the male terminal appendages,
sinensis differs from most other Habropoda by the lateral shifting and reduction of the
transverse ridges at the constriction of the 7th gastral sternite, whose diamond-shaped
apical plate is covered with very fine, short setulae; in cther details of male genitalia,
the species is a true Habropoda (Fig. 2—3). Although Popov did not mention sinensis
in his revision, he was apparently inclined to place it in Emphoropsis (see above),
a genus whose characters, so far known, are hardly in accordance with those of the
present species.

In a letter dated March 1, 1968, Dr. T. C. Maa called my attention to a recent publica-
tion by (Miss) Yen-Joo Woo (loc. cit., 1965), in which the author deals with the more
common Anthophorine species occurring in China. This account, which I have not been
able to consult, is provided with keys to genera and species, with brief descriptions of
nine species, including H. sinensis Alfken. According to Dr. Maa, it is of interest to note
that Miss Woo's article is based mainly on the collections of the Musée Heude, in
Shanghai (now known as Shanghai Station, Entomological Institute of the Academia
Sinica). The specimens of H. sinensis are topotypical and were labelled as such many
years previously by Dr. Maa himself, who therefore assumes that they were correctly
identified. He states that this bee is evidently widely distributed in the submontane
areas of China south of the Yangtze River.

In the province of Fukien, the flight season of sinensis is given by Maa as lasting
from June till October. It was repeatedly found in situations similar to those of H. imi-
tatrix spec. nov., and apparently during the same time of the year.

M. A. LIEFTINCK: Review of Asiatic Habropoda 181

ANS

am II RE
NN Ò

Sy
\ —
ANNE
S

TOS N
NA
N x
ù INN N

7
5
7
j
/
Y
/
7
a
/
4
4
tA
/

r
7
2
7
z
z
/
7
e
2
2
/

Fig. 1—3. Habropoda sinensis Alfk., & structures, Fukien; Fig. 1, exterior view of left hind tibia-

basitarsus; Fig. 2, eighth and seventh sternites, exterior view; Fig. 3, right gonoforceps and penis

valves, partial ventral view, showing appendages: intero-ventral lamella and gonostylus. — Fig. 4—7.

H. mimetica Ckll., & structures, Fukien; Fig. 4, exterior view of left hind tibia-basitarsus; Fig. 5,

eighth and seventh sternites, exterior view; Fig. 6, right gonoforceps with appendages and penis
valves, partial ventral view, with detached right gonostylus, dorsal view

182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Habropoda tumidifrons spec. nov.
Fio. 9, 12

Material examined. — Indochina: 1 9, South Viet Nam, 6 km south of
Dalat, 14—1500 m, 9.vi-7.vii.1961, N. R. Spencer coll. The specimen is the holotype
(BISH).

Female. — A more colourful species than H. sinensis, with linearly banded abdomen.

Labrum subrectangular, almost two times broader than long, widest basally (ratio
100 : 52); dark brown, basal tubercles small, smooth, rest of surface coarsely punctate,
all punctures coalescent and of large size, disk covered sparsely with light brown
semierect hairs, densest and tufty mid-apically; anterior border slightly convex, median
emargination unapparent, disk with pair of feebly raised, transverse, anteapical ridges.
Basal half of mandibles yellowish brown, for the rest black. Face evenly convex in
profile, protruding anteriorly by less than two-thirds greatest diameter of eye (63 : 100),
projecting portion of clypeus (frontal view!) distinctly broader than long (ratio 66 : 50);
almost hairless, but paraclypeal and parorbital areas covered with short appressed whitish
tomentum; raised plumose hairs on vertex and ocellar area dark brown, the long occipital
fringes and genal hairs again lighter. IOD : OOD = 60:100. Antenna brown, the
anterior faces from end of segment 4 as far as the apex, red-brown; 3 exactly twice as
long as broad at apex, length ratios of 3, 4 and 5 = 100 : 44 : 60.

Tegulae light brown, smooth, with few, very fine, superficial punctures. Punctures
covering mesonotum and scutellum more crowded and deeply impressed than those on
summit of head; gastral tergites again finer and more superficially punctate, the inter-
spaces hardly exceeding one puncture width. Basitibial plate forming an isosceles triangle
with broadly rounded apex, surface flat, finely tessellate, margins distinctly raised and
rim-like. Pubescence on dorsum and upper part of sides, as well as base of tergite 1,
rather short, moderately dense (not quite concealing surface), much as in sinensis, but
deeper in tint, ochraceous-orange, the hairs becoming paler ventrally. Broad, compact
hair-band at apex of tergite 5 and sides of 6 dark brown; posterior fringes at sternites
2—4 pale yellowish, longest at sides of 4.

Habropoda orbifrons spec. nov.
Fig. 10, 13

Material examined. — Thailand: 1 9, NW Thailand, Chiangmai, Doi Pui,
1500 m, 2.v.1958, T. C. Maa coll. The specimen is the holotype (BISH).

Very similar to H. tumidifrons. In addition to the characters given in the key, a minor
difference is found in the punctation of the abdominal tergites, the punctures being
more numerous and a little smaller than in twmidifrons. Face a little more protruding
anteriorly in proportion to greatest diameter of eye (ratio 73 : 100), and clypeus in
frontal view a trifle broader (ratio of length to breadth 50 : 70), than in tumidifrons.
Antennae as in that species, 3 a trifle longer (length: breadth = 33 : 15), length ratios
of segments 3, 4 and 5 = 100:51: 66. IOD :OOD = 70 : 100.

In both species the second submarginal cell of the fore wing is almost square, very
little broadened distally (in sirensis more markedly so). First recurrent nervure entering
second submarginal a little before distal side of the latter (in sinensis variable and
frequently interstitial).

M. A. LIEFTINCK: Review of Asiatic Habropoda 183

Undoubtedly intimately allied to tamidzfrons and easily recognized by a combination
of characters enumerated under sinensis. The male of the present species being still
unknown, nothing can be said about its internal structures, yet we may expect a form
of apical abdominal sternites and genitalia resembling those of tumidifrons and sinensis,
these organs being known only from the last-mentioned one.

Habropoda mimetica Cockerell
Biel, 15-16

Habropoda mimetica Cockerell, 1927, Amer. Mus. Novit., 274: 15 (@ Yen-ping [Fukien}, China,
Sept. 11, 1920, H. R. Caldwell) — Lieftinck, 1966, Tijdschr. v. Ent., 109: 146 (notes).

Material examined. — SE China, Fukien Prov.: 7 g (one diss., Fig. 2—6),
10 2, Shaowu distr., Kuhsienkai, 300 m, no dates, 1944 (4 g' 6 9), Shaowu City,
300 m, 18.x.1941 (1 9), no date, 1942 (3 G 3 2); Kienyang distr., Hwangkeng,
ca. 500 m, 11.x.1943 (1 2); all T. C. Maa (ML and various other collections).

Original description. — “Female — Length about 15 mm; robust, black, the legs
dark red, and the abdomen reddened at base below; clypeus entirely black, but a broadly
triangular cream-colored supraclypeal mark; antennae black, slightly stained with reddish,
the flagellum dusky ferruginous beneath; labrum with pale golden hair; face, front and
vertex with thin black hair; cheeks with white hair; thorax with bright yellowish-tawny
hair, except a very broad dark brown or nearly black band between the wings; tegulae
very dark; wings very brown; second cubital cell broad and quadrate, receiving recurrent
nervure some distance before its end; marginal cell long, not much less than half its length
beyond the third cubital cell; legs with black or brown-black hair, dark reddish on inner
side of tarsi, long and yellowish-white on under side of anterior femora, also pale on
middle legs beneath basally, anterior and middle tibiae with a little red tuft on outer
side at apex; spurs dark; abdomen densely hairy, first segment with long fulvous hair,
except at apex, where it is black; second with black, but a fulvous fringe at apex; third
and fourth very bright orange-fulvous; apex with pale chocolate hair; hair of venter
long and mainly pale chocolate. — Yen-ping, China, Sept. 11, 1920. Collected by the
Rev. H. R. Caldwell. Allied to H. turneri Ckll. (Assam) by the long marginal cell and
general appearance, but H. turneri has the black band on third abdominal segment, and
the thorax anteriorly all black. H. rowlandi Meade-Waldo (Assam) has the black band
on second segment, but no black band between the wings. Anthophora pseudobomboides
Meade-Waldo (Assam) looks superficially like H. twrneri, but has the second cubital
cell narrowed above, short appressed felt-like tomentum on second abdominal segment,
which is black at sides, etc. These insects present an extraordinary resemblance to the
species of Bombus; the present species (H. mimetica) recalling such species as B. bizo-
natus Smith and B. mendax subsp. defector Skorikov.”

A conspicuous, sturdily built species of large size, unlike any of its congeners, and
superficially resembling a short-haired bumblebee, hence the name.

Female. — Though complying with the above description, some additional peculiarities
are worth mentioning. Proboscis relatively short, reaching fore coxae in repose: galea
not longer than clypeus and labrum united. Last segment of labial palpus shorter than
penultimate joint. Labrum brownish black, all punctures large, coarse and coalescent, disk
bounded on either side by a low, slightly outcurved ridge placed in the long axis;
thickened semicircular border of emargination obliquely cut off, the brushy median

184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Fig. 7—10. Frontal view of heads of Habropoda, semi-diagrammatic; Fig. 7—8, H. sinensis Alfk.,

Fukien; Fig. 9, H. tumidifrons spec. nov., holotype, S. Viet Nam; Fig. 10, H. orbifrons spec. nov.,

holotype, Thailand. — Fig. 11—15. Pygidial plates of © Habropoda, dorsal view; Fig. 11, H. sinen-

sis Alfk., Fukien; Fig. 12, H. tumidifrons spec. nov., holotype, S. Viet Nam; Fig. 13, H. orbifrons

spec. nov., holotype, Thailand; Fig. 14, H. disconota spec. nov. (apical portion, more enlarged),

N. Viet Nam; Fig. 15, H. mimetica Ckll., Fukien. — Fig. 16. Frontal view of head of 6 H. mime-
tica Ckll., Fukien

tuft golden brown. Tufts of long, raised and mostly dark hairs before and behind
anterior ocellus, and all along upper occipital border, the fringe behind the latter and
on genal area, pale yellowish to white. IOD : OOD = 69 : 100. Antennal scape slender,
subcylindrical, not flattened anteriorly, shortly pilose, only little longer than segment 3,
which is almost as long as its width at apex (100 : 34.6), end equalling in length the
next three (4—6) combined; ratio of 3, 4 and 5 = 100 : 25 : 30.8; colour blackish .
brown, 4—12 ochraceous anteriorly. Body punctation extremely dense, except smooth
and glossy mid-propodeal triangle. Raised silky hairs covering thoracic segments com-
pletely hiding integument. Tegulae brownish black, very shiny, with few minute super-
ficial punctures. Wing membrane light smoky yellow, apical border hardly obscured.
Venation as described for sinensis, the second recurrent nervure never quite interstitial.
Basitibial plate broadly oval, with slightly raised margin and overlaid basally with golden,
matted hairs originating from apex of femur. Tarsal arolia absent, or, when vestigial,
then flat, subcordate.

Male (hitherto undescribed). — Mouth parts retracted, galea reddish brown, shorter
than in female, not reaching fore coxae. Mandibles black, only the mid portion chestnut.
Labrum jet-black, the broad polished median line variable in breadth, complete or
confined to its distal half. Face even less protuberant than in female. Chrome patch
occupying most of clypeus trapezoidal in shape, the tentorial pits vestigial, not or only

M. A. LIEFTINCK: Review of Asiatic Habropoda 185

slightly obscured. Antenna brownish black, flagellar segments as in female; scape some-
what flattened and broader, its antero-lateral face fringed with long, raised yellowish
hairs; segment 3 markedly shorter than scape. Eyes considerably larger and more convex
than ín female, their shortest distance (at level of antennae) and own diameter, in
frontal view, in the ratio of 30 : 25. Body sculpture as in the opposite sex, pubescence
of the same length, but much paler and showing a different pattern: dark band across
mesonotum as broad as in female, but not at all defined, most fine plumose hairs between
the wings tipped with grey-brown at the transition into the pale orange-yellow hairs
covering remaining parts of the dorsum; thorax greyish white underneath. Wing mem-
brane as in the female; second submarginal cell frequently almost square, and position
of second recurrent nervure also variable, either interstitial, or (more often) entering
the cell just before distal side of the latter. Abdominal pubescence as described in the
key, the black band on tergite 2, so conspicuous in the female, replaced by yellow, disk
in very fresh examples only more distinctly brown, or with admixture of darkish hairs,
when looked at from the side.

Habropoda disconota spec. nov.
Fig. 14

Material examined. — Indochina: 1 9, North Viet Nam [Tonkin], environs
of Hanoi, no dates, Thank leg., ex coll. B. Tkalcu. The specimen is the holotype (ML).

For a characterization of this bee, see the key to the females. Undoubtedly intimately
related to mimetica, approaching the latter so closely in general appearance and major
external features that no further description is needed. Originally I thought this to be
only a geographical subspecies of mimetica, but, apart from the dissimilar colour design
noted, I am now convinced that the remaining characters of this unique specimen are
sufficiently distinctive to warrant specific recognition. One of the really noteworthy
features common to both is found in the clypeus, which does not protrude strongly
from the face, but arises gradually from the plane of supraclypeal and paraclypeal areas.
Unfortunately, the male of disconota still remains unknown.

Habropoda turneri Cockerell

Habropoda turneri Cockerell, 1909, Entomologist, 42: 308 (9 Shillong, Assam) — Lieftinck,
1966, Tijdschr. v. Ent., 109: 145—146 (notes on 2); 1972, Tijdschr. v. Ent., 115: 281 (host-
parasite relationship).

Type material examined. — NE India: 1 9 (holotype), “Assam, Shillong, 9.03”
(written), labelled “type” and “Habropoda turneri Ckll. Type”, in Cockerell’s hand-
writing (general coll., BM).

Further material examined. — NE India: 19 (topotypical), same labels as
holotype (BM); 1 & (diss., Fig. 17—20, first described J'), “Shillong, 9.03‘ (written),
“Assam, R. Turner 1905—125” (print) (BM); 1 &, “Shillong, 8.03” (written),
“Assam, R. Turner 1910—225” (print) (ML); 1 9, “Shillong, 8.03” (written) (ML).
1 & (head and proboscis detached, abdomen partly missing), “Coll. Friese” (print),
“Khasi Hills, 1.9.95” (written, blue disk), “Anthophora bremoides Fr.” [nom. nud.},
det. Friese 1913 (MNB).

A rather large, handsomely coloured species, easily distinguished from others by the

186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

conspicuous variegated colour-pattern, dark legs and fuliginous wings. For the sake of
completeness, the original description is here quoted in full:

“Q. Length about 17 mm., anterior wing 11; robust, but with the abdomen much
longer in proportion to its breadth than in H. zonatula, Sm.; black, with the pubescence
of the head, and thorax above as far back as the level of the hind wings, black; posterior
to this, beginning abruptly, the pubescence is sulphur-yellow, and the same, very dense,
covers the first two segments of the abdomen; the third segment has the hair short,
dense and coal-black; the fourth and fifth have it reddish, more or less black at sides;
there is a little pale tomentum at sides of face, and the lower part of the cheeks is
covered with long white hair; a peculiar feature is a small patch of appressed white hair
just above (a little mesad of) each antenna, surrounded on all sides by erect long black
hair; the pleura is covered with long white hair; hair of legs mainly black, but some long
white hair on anterior femora beneath; brush at end of hind basitarsus orange-fulvous.
Clypeus prominently densely punctured, but the punctures irregular and largely in
grooves; antennae black, third joint a little longer than 4 + 5; tegulae black; wings
fuliginous, third s.m. broader than second; marginal cell long; hind tibiae broad and
flat, not produced at end, the scopa coarse and dense.

Hab. Shillong, Assam.

A very distinct species, not closely resembling any of the four (H. montana, Rad.,
H. magrettii, Bingh., H. moelleri, Bingh., H. fulvipes, Cam.) known from India. There
is a certain general resemblance to H. tarsata, but that has the hair of the thorax entirely
orange-fulvous above.”

(It may be noted in passing, that all species compared by Cockerell with twrnert,
except montana (= radoszkowskii), have since been transferred to Elaphropoda Lieft.).

Female. — Maxillary palpi slender, 1st segment short, only twice as long as broad,
the 2nd longest, and from 3 on gradually diminishing in length, the whole palpus
shaped rather like Elaphropoda (see Lieftinck, 1966: 126 fig. 18). Proboscis light
brown, much longer than in H. mimetica, measuring 10 mm when fully extended, galea
reaching end of hind coxae in repose. Dark pubescence covering anterior portion of
thorax extending below wings about as far down as the scrobal groove, the large patch
below this silvery white; coxae and femora predominantly dark-haired, as also the
abdominal sternal fringes, which are of moderate length, most conspicuous at sides of
4 and 5. Basitibial plate blackish, broadly oval, surface dull, quite flat and not rimmed,
the apex rather protuberant. Tibial spurs light brown. Inner ramus of mid and hind
tarsal claws only half as long as outer and more strongly curved than the latter. Wing
membrane light brown; marginal cell of fore wing long and attenuated, relatively a little
longer than in male (ratio 100 : 75.5); second submarginal cell of the usual, almost
square, shape, the recurrent vein entering the cell a little before its distal side. Pygidial
plate, Fig. 21.

Male (hitherto undescribed). — Unidentified specimens of this sex in the British
Museum (Nat. Hist.) collection turned up unexpectedly in drawers containing various
other anthophorines, after publication of my 1966 paper (loc. cit.).

The principal characters are those given in the descriptive key. Very similar to the
female, but size a little smaller and all colours less bright, the dark band on gastral
tergites 2—3 less definite, anteriorly as well as posteriorly. Proboscis of the same great
length (9 mm approx., small specimen), exceeding length of abdomen. White pubescent
patch of finely plumose hairs behind antennae directed anterad, occupying the depressed

M. A. LIEFTINCK: Review of Asiatic Habropoda 187

S
ST

—
TTT II
+ a
IA

SMA

IN
NA

DINIEREN

SEN as,
SS
NESS
WSS

A N
ANS

Fig. 17—21. Habropoda turneri Ckll., structures, Shillong; Fig. 17, exterior view of & left hind
tibia-basitarsus; Fig. 18, ventral view of & seventh tergite; Fig. 19, eighth and seventh sternites of
4, exterior view; Fig. 20, right gonoforceps with appendages and penis valves of &, partial ventral
view; Fig. 21, pygidial plate of 9, dorsal view

facial foveae on either side and confluent with longer white hairs in the middle of the
supraclypeal area. Tegulae brown, surface shining, clothed with longish, darker brown
hairs hiding most of the surface. Hair on outer face of hind tibia and basitarsus all
black, much less condensed than in female, not hiding the surface, which is rather
shining and evenly punctate; tarsal distalia brown, with yellowish brown hairs, last
segment and claws basally, ferruginous tipped with black. Apex of tergite 7 excised,
dorsal surface clothed densely with long, bristle-like hairs (Fig. 18). Sternites 7 and 8
of characteristic shape (Fig. 19—20). Intero-ventral process at apex of gonocoxite placed
in a vertical plane: when viewed from behind in the form of a broad, bluntly rounded,

188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

almost bare, oval plate, which is directed inward and hollowed out externally; owing
to its transverse position, the plate in ventral view appears to be slenderly lanceolate
(Fig. 20); gonostylus much longer, base slender, ribbon-like, the flattened distal portion
gradually distended, its apex fringed with long, stiff bristles.

With the discovery of the males of both mimetica and turneri, it became quite evident
that the two are not at all nearly allied, as had been held previously by Cockerell and
myself. These statements were based on the long marginal cell and reduced tarsal arolia,
characters common to both. In point of fact the resemblance between them is merely
superficial, due to a somewhat similar pubescent colour-design. The pronounced sexual
dimorphism of mimetica is non-apparent in turneri, which is more slenderly built and
has a differently shaped head, the legs and hidden sternites of the male being also
very unlike those of mimetica. In general appearance, and on comparing the structural
features of the males, turneri approaches rowlandi, discussed hereafter, much more
closely, all facts suggesting real relationship.

In a recent publication (Lieftinck, 1972), I suggested the Indian melectine Proto-
melissa tricolor m., also from Shillong, as a possible parasite of H. turneri.

Habropoda rowlandi Meade-Waldo

Anthophora (Habropoda) rowlandi Meade-Waldo, 1914, Ann. Mag. Nat. Hist. (8) 13: 45 (key),
50—51 (2 & Shillong, Assam, viii.1903, R. E. Turner).
Habropoda rowlandi: Lieftinck, 1966, Tijdschr. v. Ent., 109: 146 (brief notes on both sexes).

Type material examined — NE India: 1 9 (holotype, no. 637, selected by
Meade-Waldo), “Assam, Shillong, 9.03, R. E. Turner, 1910—225” (BM).

Further material examined. — NE India:3 g,3 © (first described &, diss.,
Fig. 23—25, BM), “Assam, Shillong, 8.03” (written), “Assam R. E. Turner, 1910—
225”, and “Turner Coll. 1912—111” (print), Q with additional label “Habropoda 9 ”
(general coll, BM and ML); 3 g, 1 9, “Shillong, 8.03, Assam” (written), R. Turner
1905—125, one with label “Habropoda 2” (BM).

Apparently based on several specimens of either sex, but with the exception of the
holotype and one male standing over a drawer label row/andi, all were unidentified.

Note. — A number of “rowlandt”’ in the Brit. Mus. (Nat. Hist.) collection with
typewritten labels “Simla, 8.98” and “9.98 Bingham coll”, and placed as rowlandi by
Meade-Waldo, are not that species but a mixture of apatelia spec. nov., apostasia spec.
nov., and deiopea Cam., discussed hereafter.

My statement (1966 : 146) that the type of this species is not a true Habropoda,
is incorrect, in so far as it was based on characters which I now consider are of no
generic importance. H.rowlandi is, I think, most closely related to twrneri, a species
whose male had not been recognized at the time, but which is described in the previous
pages.

So far known only from Meade-Waldo’s nondescript diagnosis. The specimens now
before me form a homogeneous series showing little or no variation.

The two sexes are characterized anew in the keys, which can be supplemented as
follows. Stature slenderer and of smaller size than twrneri, from which it is at once
distinguished, among other characters, by the more uniformly coloured body pubescence,

M. A. LIEFTINCK: Review of Asiatic Habropoda 189

Y 4

5 7
7
y
7
7
LA
4
?
4
4

Fig. 22—26. Habropoda rowlandi Meade-Waldo, structures, Shillong; Fig. 22, exterior view of &

left hind tibia-basitarsus; Fig. 23, ventral view of & seventh tergite; Fig. 24, eighth and seventh

sternites of 4, exterior view; Fig. 25, apical portion of genital capsule, showing gonoforceps with
appendages and penis valves of 6, ventral view; Fig. 26, pygidial plate of 9, dorsal view

lighter brown and palely pubescent legs, and also by the presence of a well-developed
tarsal arolia and much lighter wings.

Male and female. — Proboscis of the same great length as /urneri, galea reaching
as far back in repose as the mid coxae. Labrum broader and more angulated laterally,
lacking the dark crescentic ante-apical ridge of turneri, and with the discal hairs (as well
as the mid-apical and lateral fringes) much longer than in that species; sides of clypeus
with light marginal hairs also more conspicuous and longer, especially so in male. Malar

190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

area distinctly longer than in turneri. Legs much lighter, the tibiae and tarsi more
reddish brown, clothed all over with brown hairs, those covering mid and hind tibiae
and basitarsi light golden brown. Tibial spurs golden yellow; tarsal claws with distinct,
slender, laterally compressed, arolia, otherwise shaped as in fzrneri. Wing membrane
pale smoky yellow. Vestiture of abdomen composed of a matted layer of plumose
pubescence covering the tergites posteriorly, and much longer, raised and finely branched
hairs more sparsely disseminated over the whole surface, all discal hairs shorter than
in turneri.

Male. — Apical fringes of tergites 6—7 and sternite 6 with distinct golden gloss.
Hidden sternites 7—8 (Fig. 24) considerably more expanded than in ##rneri, but strong
transverse ridges at base of apical plate and abundant short microsetae similarly arranged.
Structure of genitalia likewise resembling that of turneri, the intero-ventral lamella at
apex of gonocoxite and gonostylus both broader than in that species (Fig. 25).

Female. — Basitibial plate dark brown, shaped similarly to mimetica, with the same
golden brown femoral tuft overlying its base, thus differing from turneri. Apical fringes
of tergites 5—6 with golden shine, as in the male. Pygidial plate, Fig. 26.

A peculiarity of the venation not mentioned earlier, is found in the shape of the sub-
marginal cells of the fore wing. Considerable variation exists in the proportional sizes
of the second and third submarginal cells, the latter frequently being scarcely broader
than the second. In turneri and most other species, the third s.m. is much larger than
the second, the costal and anal sides of the latter being markedly shorter than those
of the third cell. In most rowlandi, on the other hand, the third is only little larger
than the second, in several males the reverse condition existing as to the lengths of the
costal and anal sides of these cells.

Habropoda tainanicola Strand
Fig. 27—37

Anthophora (Habropoda) tainanicola Strand, 1913, Suppl. Ent., 2: 51—52 (@ Tainan & Hoozan,
Formosa).

Anthophora bucconis Friese: Strand, 1913, Suppl. Ent, 2: 51 (4 ® Formosa, addit. descr.,
partim!).

Emphoropsis spec.: Michener, 1944, Bull. Amer. Mus. Nat. Hist, 82: 286 (& Formosa).

Habropoda tainanicola: Lieftinck, 1966, Tijdschr. v. Ent., 109: 146 (notes).

Type material examined. — Taiwan (Formosa): 1 9, “Cotype” (print on yellow
disk), “Tainan, Formosa, 1911, H. Sauter/7.IV” (print), “Anthophora tainanicola m.”,
in H. Strand’s writing/“Cotype” (print on dark red), “H. tainanicola Strand”, det.
M. A. Lieftinck 1967 (BM).

Further material examined. — Taiwan (Formosa): 1 9, “Formosa, Takao, 3.
1908, Sauter/Anthophora pyropyga Friese 1915 [nom.nud.} Type” (print on dark
red), “tainanicola Str., det. H. Hedicke 1942, cpd. with cotype by M. A. Lieftinck 1970”
(MNB). 3 & (first 4, one diss., Fig. 28—31), 6 2 (2 9 light 49 dark form),
Taiwan, Neihu nr. Taipei, 250 m, iv-v.1972, at nesting site; 15 & (1 diss.), 10 ©
(4 9 light, 3 Q dark, 3 9 intermed. form), same locality, 20—25.iii.1973, 1—2.iv.
1973; all T. C. Maa (ML, etc.). 1 & (diss.), Formosa, Urai, i.iv.1932, J. L. Gressitt,
“Habropoda spec.”, det. C. D. Michener (UKL).

This remarkable species stands rather apart from its Asiatic congeners Although both

M. A. LIEFTINCK: Review of Asiatic Habropoda 191

sexes bear a striking superficial resemblance to H. bucconis and the nearly related
imitatrix spec. nov., tainanicola can be distinguished from these by a number of structural
peculiarities described in the key. The most striking differences are found in the abdo-
minal sternites of the male and pygidial plate of the female; H. fainanicola also deviates
from the two regional species just mentioned by the colour and sculpture of the face,
the male moreover by the placement of the ocelli, the length ratios of the proximal
antennal segments, and the long hairs fringing the outer ridges of the hind legs. When
good series are available for comparison, it will be seen that tainanicola averages a little
smaller in size, the insect at the same time appearing somewhat more slenderly built,
| than bucconis and imitatrix.

There appear to be two distinct geographical subspecies: one, the nominotypical
tainanicola, described from the island of Taiwan (Formosa), and a second, matella
subspec. nov., from the opposite Chinese continent. In respect of size, general morpho-
logy and sculpture, the two races are practically identical, males in addition being
inseparable as regards shape and armature of the hidden sternal plates and genital
organs. In spite of this, males can be recognized fairly easily by a different pubescent
colour pattern, which appears to be constant in both. The females are dichromatic, there
being two colour forms of either subspecies, one (isochromatic) resembling the male,
and a second (heterochromatic), almost black-haired, form with a “red-típped” abdomen.
The latter is the one described by Strand as tainanicola. Specimens intermediate in colour
do occur in populations of both subspecies.

H. tainanicola tainanicola Strand
Fig. 27—32

Here follows first the original description:

“9 9 Q: Tainan 7. April, Hoozan April. 3

Die Art muss wohl für eine Habropoda gehalten werden, wenn auch das Geäder
nicht konstant wie bei dieser Gattung oder Untergattung ist: bei dem einen Exemplar
ist die erste rekurrente Ader interstitial, bei den übrigen mündet sie in die zweite
Cubitalzelle ein, aber allerdings ganz nahe der zweiten Cubitalquerader. Labrum mit
hellem langen Haarschopf. Die Art ahnelt H. Radoszkowski D.T. (= montana Rad.),
aber Clypeus ist einfarbig schwarz, die Behaarung des Thorax und der Basalhälfte des
Abdomens ist schwarz usw.

Tegument schwarz, an den Fiihlern ist die Spitze des Schaftes und die Unterseite des
Flagellums, abgesehen vom Endglied, gerötet, an den Beinen sind die Spitze der Femoren,
die ganzen Tarsen und wenigstens die Spitze der Tibien, am II. und III. Paar die ganzen
Tibien gerötet. Behaarung schwarz, rotgelb (feuerrot) aber an den Segmenten 4—6,
wenigstens der Hälfte des Segmentes 3, an allen Metatarsen und Tarsen, an den ganzen
Tibien 3 und an der Spitze der Tibien 1—2; letztere beiden haben auch oben mehr
oder weniger rotgelbe, mit schwarz gemischte Behaarung, und blass rötlichgelbe Be-
haarung findet sich an der Unterseite des Körpers, den Schläfen und an den Femoren,
hell rötlichgelbe auf den Hinterrindern der Bauchsegmente, dem Labrum und (mit
schwarz gemischt) auf dem Clypeus. Tegulae schwarz. Die Flügel sind gelblich ange-
raucht, stark schimmernd, mit braunschwarzem Geäder, das Saumfeld (bis an die
Zellen) am dunkelsten. Clypeus stark vorstehend, vorn in der unteren Hälfte flach, der
dieser Fläche entsprechende Teil des Vorderrandes gerade und ganz schwach erhöht,

192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

rs!
AST
N
vs 4,
(N u
È 4
\ 7,
È ‘7
N 4a
d 2
È 2 po,
N 7
< te
Ò >=
= ves

fd
=

ee VAGLIA DA

Fig. 27—32. Habropoda t. tainanicola Strand, structures, Taiwan; Fig. 27, exterior view of & left

hind tibia-basitarsus; Fig. 28, ventral view of & seventh tergite; Fig. 29, exterior view of & sixth

sternite; Fig. 30, eighth and seventh sternites of 4, exterior view; Fig. 31, left gonoforceps and
penis valves of 4, partial ventral view; Fig. 32, pygidial plate of 9, dorsal view

der ganze Clypeus etwas glänzend, und zwar vorn mitten am deutlichsten, mit ganz
seichten und nicht dichtstehenden kleinen Grübchen. Der Scheitel stärker glänzend,
zwischen Ozellen und Augen seicht ausgehöhlt, glatt, das Ozellenfeld matt. Die Wangen
schmal, glänzend, glatt, jedoch mit einzelnen ganz feinen Pünktchen. Mesonotum,
soweit durch die ziemlich dichte Behaarung erkennbar, ist etwas glänzend, fein und
spärlich punktiert. Das zweite Geisselglied ist jedenfalls nicht länger als die drei folgen-

M. A. LIEFTINCK: Review of Asiatic Habropoda 193

den zusammen. Die dritte Cubitalquerader ist gleichmässig saumwärts stark konvex
gekrümmt, die erste ist umgekehrt und zwar ungleichmässig und fast unmerklich ge-
bogen. — Körperlänge 14 mm. Flügellänge 10,5 mm. Breite des Abdomens ca. 6 mm.”

This good description complements the female characters given in the specific key
and the more detailed racial distinctions given below.

Male. — Vestiture of body throughout bright orange-chrome to orange-rufous, the
long pubescence on supraclypeal area, mandibles, genal area, ventral surface of body
and bases of fore legs, lighter, pale orange-yellow; dark brown raised hairs only on
paraclypeal area, but erect tufts on summit of head in front of ocelli and at occipital
border also obscured; occasionally some brown-tipped hairs interspersed on mesonotum
anteriorly. Broad membranous posterior margins of tergites 1—6 yellowish, well visible
beneath the pubescence. Wing membrane stained with pale orange-yellow, outer borders
of both fore and hind wing broadly and distinctly obscured.

Female. — (a), typical (dark) colour form: see original description. Apart from
the “cotype”, four out of our series of nine dark females have the abdomen as described
by Strand, whereas in five others tergites 3 and part of 2 are likewise red. (b), light
colour form: the remaining seven females are isochromatic, coloured similarly to the
male though having more grey-brown hairs intermixed on the thoracic dorsum; the
other three are intermediate, differing from the rest by having most of the thorax
clothed with dark rufous-brown hairs and a vivid orange-rufous abdomen, more deeply
so in tint than in the male. The legs have retained the same bright colour throughout
our series of females.

The proboscis is almost as long as in H. twrneri and rowlandi, the galea measures
4.2—4.6 mm, and the tongue when fully extended, reaches as far as the apex of the
trochanters of hind legs. The basitibial plate of the female is rimmed, shaped similarly to
that described for mimetica and rowlandi, with the same conspicuously golden basal tuft
at apex of femur as seen in rowlandi. Venation light brown; shape of submarginal cells
variable, costal side of the 2nd always a little shorter than anal side, the 3rd cell dis-
tinctly longer than 2nd; recurrent nervure either interstitial, or received a little before
distal side of 2nd submarginal.

Strand only described the female, from ‘‘Sauter’s Formosa Ausbeute”, a number of
females —and possibly also light-coloured males — having been confounded by him
with H. bucconis in the same publication; see under that species.

H. tainanicola maiella subspec. nov.
Fig. 33—37

Habropoda tainanicola: Cockerell, 1931, Amer. Mus. Novit., 480: 7 (2 Foochow distr.).

Material examined. — SE China, Fukien Prov.: 11 4 (2 diss.), 52 9, Shaowu
distr., Tachulan, 1000 m, iv-v.1934, 1942—43 ( S diss., 21—27.iv.1943 and 11.iv.1943,
on white labiate); 3 G', 14 9, Chungan distr., Tsilichao, 1000 m, 4.iv.1943 (2 g'),
Sanchiang, 600 m, 30.iv.1928 (J') and iv.1942; 12 9, Chungan distr., Bohea Hills,
600 m, iv.1940; 3 9, Kienyang distr., Hwangkeng, 22.iv.1943; 1 9, Central Fukien,
Changting, 27.iv.1943, at Citrus; all T. C. Maa (ML, etc.). 3 9, Fukien, Kushan
below Foochow, 400 m, 8.iv.1935, and Buang Kri, north of Foochow, 26.iv.1935 (inter-
mediate colour form), E. Suenson (BISH). Holotype 4 and first described 9 : Shaowu,

194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Il if
IN
A 7
\ /
{ vf
U 7
! /
N no
Ÿ iy
N HA
GTA
N nF
N Teo)
È DAN,
VA
ne ; +4
Se 5
ui 7
RS wT

Fig. 33—37. Habropoda tainanicola maiella subspec. nov., & structures, Fukien; Fig. 33, exterior

view of left hind tibia-basitarsus; Fig. 34, ventral view of seventh tergite; Fig. 35, exterior view of

sixth sternite; Fig. 36, eighth and seventh sternites, exterior view; Fig. 37, left gonoforceps and
penis valves, partial ventral view

Tachulan, 21—27.iv.1943 (Lg diss.), and same loc., 2.vi.1942 (9) (ML); paratypes
in various other collections.

This is obviously the same bee as the one which Cockerell characterized briefly as
follows: “The single female sent is in very bad condition, but it appears to belong to
Strand’s species, described from Formosa. It is very distinct by the entirely black face;
the dark reddish labrum keeled down the middle and sharply bidentate at end; the large

M. A. LIEFTINCK: Review of Asiatic Habropoda 195

red area on basal part of mandibles; brownish black hair of head and thorax (but
cheeks below with long gray hair); dark tegulae; yellowish-hyaline wings; hind tibiae
and tarsi covered with very bright ferruginous hair; apex of abdomen with red hair.
If this is at all different from the Formosan insect, the fact can only be ascertained
by direct comparison of good specimens.”

Male. — Vestiture more sallow and generally darker than in ¢. Zainanicola; long
pubescence on labrum, supraclypeal area, genal surface and fore femora almost pure
silvery white; summit of head, mesonotum and scutellum a mixture of grey-brown, sides
below wings dark brown, pleurae pale brownish fading to almost white ventrally. Legs
light to dark brown, the dorsal fringes of hind tibia and basitarsus variable, frequently
almost black. Wing membrane much as in the insular race but outer border of wings
less obscured. Broad membranous posterior margins of tergites 1—6 pale ochraceous;
pubescence of 1 greyish yellow to pale orangish gradually darkening to grey-brown on
succeeding segments, 4—5 again lighter with orangish hue, 6 and 7 progressively more
vividly orange-yellow, the lateral tufts at tergites 4—6, tomentum of sternites 5—6,
as well as the dense apical fringe of 7, light golden yellow in fresh specimens.

Female. — (a), typical (dark) colour form: very similar to nominotypical examples,
but differing constantly in that the dark hair covering thorax and abdomen, besides
being dark brownish black instead of deep black, is thinner, leaving visible more of
the underlying surface, especially that of the abdomen. It differs further by the reduction
of light colour on the abdomen, the black extending as far as a little before the apex
of tergite 4, the posterior fringe of which is light orange-yellow instead of rufous, only
the dense hair covering tergite 5 acquiring a golden-rufous tint. Long bristles fringing
tergite 6, on either side of the pygidial plate, glistening golden brown. (b), light colour
form: much lighter and less uniformly coloured than the lightest individuals of
t. tainanicola. Eight females (about 10% of the total) are isochromatic, though differing
from the male in that they are more densely hairy, the pelage on the whole being less
sallow, brownish yellow, lightest on the scutellum, the first and penultimate abdominal
segments, the intermediate ones acquiring a more greyish brown tint, while 5 and 6 are
again orange-rufous, these colours on all parts shading one into the other. About 15%
of the total number of females are more or less intermediate in this respect.

Attention should be drawn to the accompanying Fig. 27—32 and 33—37, of the
two races of fainanicola, which were prepared only from a single individual of each
taken at random. The apparent differences brought about in these drawings give no
answer to the amount of variation which may be observed when good series are com-
pared. So much is certain, that the form of tergite 7 as well as that of the hidden
sternites seems to vary somewhat in both subspecies, these structures being, in fact, nearly
identical in shape. —

As pointed out above, the matching of the sexes has long remained a matter of
uncertainty and, until recently, led to wrong conclusions (see below). The discovery of a
very similar subspecies in Fukien and simultaneous occurrence of its sexes in various
places on the Chinese continent, has removed all doubt about the correctness of their as-
sociation. The fact that the two races were collected in the same season already indicated
a close relationship. However, conclusive evidence could be obtained from a continued
investigation of the nesting habits of typical tainanicola in Taiwan, described hereafter.
A few years ago (Lieftinck, 1972 : 277) I have reported already on these biological
observations, which were carried out by T. C. Maa. In the spring of 1970, and again

196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

in 1971, two small colonies were discovered, clusters of cells being established in sandy
soil under overhanging rocks on a road-side slope, near the observer’s residence at Neihu
(near Taipei). At that time individuals of two kinds of differently coloured females
were observed, which occurred together with a single kind of male. The dark females
and pale brown males were identified as H. tainanicola, a species then still known only
from the female. The light colour form of the latter — though closely resembling
_ bucconis — was supposed to represent an undescribed species of which the male had
still to be discovered. The fact that the female forms inhabiting the “bee village” were
so utterly unlike, led to the erroneous assumption that a mixed colony existed of two
species, and that the male’s flight season of one of the occupants had already been over
at the start of the observations. In order to throw more light upon this question, Dr. Maa
continued his investigations the next year, re-visiting the colonies almost daily, from
the 10th of March onward till the end of April, with about one week’s interruption.
One of the nesting sites had been destroyed the year before, but the second was still left
intact. On the 10th of March, the first humming sound of a Habropoda male was heard
near the exposed entrance holes of the cell-cluster. Two days after, about 10 males (but
no females!) were seen at the site, while a small series of both sexes could be procured
from the 12th March until the 4th April, when the last male was captured, females of
all colour forms then being still on the wing. This evidently indicated that the flight
season had almost come to an end.

A careful examination of the entire series of specimens revealed beyond doubt that
all occupants of the nesting site belonged to a single species. The females are variable,
exhibiting a marked polychromatic colour design, whereas all males are of the same
uniform yellow-brown colour.

The species was found by Dr. Maa nowhere else in Taiwan, nor did he come across
H. bucconis in any part of the island. This is probably a much scarcer bee, which has
a different flight season (see under that species). The only flowers visited by tainanicola
were two species of native Rbododendron and an introduced species of Passiflora.

Habropoda apostasia spec. nov.
Fig. 38—41

Material examined. — Nepal:1 & (holotype, diss.), E Nepal, Taplejung [Tap-
lang Jong] Distr., Sangu, ca. 6200’, mixed vegetation by stream in gully, ix-x.1961,
R. L. Coe coll., Brit. Mus. Fast Nepal Exp. 1961-62 (BM).

A more easily recognized species than the other inconspicuous and small-sized mem-
bers occurring in the Himalayan region.

The present male is unique and best characterized by the slender, unmodified
form of its legs, more especially of the posterior pair. As described in the key, the
outer face of the hind basitarsus is almost flat and lustreless, very finely tessellate and
evenly covered with short, decumbent pale hairs, the pile fringing the dorsal and ventral
ridges being only little longer than the rest and, like these, of a pale yellow colour;
the inner face is closely punctate and clothed with the usual dense brush of much longer,
stiff, golden-brown hairs, which are directed apicad. It follows, that the hind basitarsus
of this species shows no expansion of its lower part, and hence entirely lacks the hairless
area, or “sole”, which is so conspicuously present in several similarly-looking species.

Proboscis retracted, but galea relatively long, almost 3.5 mm, reaching well beyond
fore coxae. Labrum, the brown basal tubercles excepted, black, the converging ante-

M. A. LIEFTINCK: Review of Asiatic Habropoda 197

-
CER

7
er me ene

i
il

Fig. 38—41. Habropoda apostasia spec. nov., 4 structures, holotype Nepal; Fig. 38, exterior view
of left hind tibia-basitarsus; Fig. 39, ventral view of seventh tergite; Fig. 40, seventh and eighth
sternites, exterior view; Fig. 41, left gonoforceps and penis valves, partial ventral view

apical ridges smooth, somewhat upturned, each ending in a low tubercle just before and
above the front margin, which is almost straight; disk as described, the pale hairs
covering it not very long, free border with short dense fringe of ferruginous hairs, the
median tuft longest. Clypeus for the greater part light yellow, surface evenly convex,
dullish, superficially, rather irregularly punctate, more shining only at apex. Antenna
dark brown, anterior face of segment 3 only with vestigial basal and apical orangish
spots, but remaining segments all brownish orange anteriorly. Long pubescence of genal
area, ventral parts of thorax and basal segments of legs, white. Other characters as
described in the key. Hidden sternal plates (Fig. 40) resembling those of sutepensis
most closely; basal portion of sternite 7 relatively broad, and transverse sclerotised ridges
reduced in size. Intero-ventral lamella at apex of gonocoxite placed transversely (Fig.
41), considerably expanded, broadest about halfway length, outer margin — when
viewed from the inside — straight, but much swollen and folded in distally, the marginal
rim thus formed bituberculate (not shown in Fig. 41). Gonostylus slightly exceeding
lamella, narrow and almost straight, the apex slenderly club-shaped in side view.

Female unknown.

This little species was collected in Nepal along with individuals of radoszkowskii,
pelmata and apatelia, all members of one group. By the clypeal marks and the yellow-
spotted antennal scape, it resembles the two first mentioned species. However, the very
different shape of the hind leg as well as the configuration of the hidden sternal plates,
are unique features which preclude the possibility of a close affinity with these. In fact,
apostasia is probably most nearly related with H. sutepensis, described below.

198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Habropoda sutepensis Cockerell
Fig. 42—45

Habropoda sutepensis Cockerell, 1929, Ann. Mag. Nat. Hist. (10) 4: 132—133 (4 Siam, Doi
Sutep, Feb. 9, 1928); Cockerell, 1930, Ann. Mag. Nat. Hist. (10) 5: 163 (® Siam, same loc.). —
Lieftinck, 1966, Tijdschr. v. Ent, 109: 146 (note on types); 1972, Tijdschr. v. Ent., 115: 257 & 281
(generic assignment and host-parasite relationship).

Type material examined. — Thailand (Siam): 1 G (holotype), “Doi Sutep,
Siam, on summit, Feb. 9, 1928, T. D. A. Cockerell’ (BM, no. 652).

Further material examined. — Thailand (Siam): 2 9, “Doi Sutep, Siam, Feb.
1928 T. D. A. Cockerell’ (BM); 1 g', “Doi Sutep, Siam, Feb. 1928, H. sutepensis
Ckll.”, det. T. D. A. Cockerell (MNB); 2 & (one diss., Fig. 43—45), “Doi Sutep,
Siam, Feb. 1928” (written), “T. D. A. Cockerell coll.” (print), “W. P. Cockerell &
A. Mackie collrs.” (written), “Pres. by Imp. Bur. Ent. Brit. Mus. 1929—570” (print),
“Habropoda sutepensis Ckll.” (in Cockerell’s handwriting) (BM & ML).

I believe to have shown in the key that, here again, we meet with a species showing
characters departing from those used for establishing the generic diagnosis.

Since briefly reporting on H. sutepensis, I have studied (1966, 1972) the morphology
of the male more carefully and arrived at the conclusion that it can best remain merged
in Habropoda along with all other species here treated. The present conception, which
holds good for a number of other aberrant members as well, is briefly explained in the
introduction.

Male sutepensis runs out in the key to near H. apostasia spec. nov., from which it is,
however, immediately distinguished by the totally different shape of the hind legs and
much longer abdominal pubescence. The hidden sternites are rather similar in the two,
but here the resemblance ends. In fact, the long malar area, unmodified — yet con-
spicuously hairy — hind basitarsus, in combination with the elongate form of the 7th
abdominal tergite, are characters peculiar to swtepensis. On the other hand, the non-
specialized long hair dress of the exposed sternites, as well as the face marks and body
punctation, definitely link the species with bucconis, so that I assume the latter to be its
nearest ally.

The key characters of the male can be supplemented briefly by the next few particulars.

Male. — Proboscis retracted, galea of moderate length, 3.5 mm approx. reaching
to end of fore coxae in repose. Clypeus ivory, basal portion with feebly indicated median
ridge, this part of the surface dull, finely, superficially and sparsely punctate, the
anterior portion slightly declivous and triangular in outline, its surface impunctate and
shining; whole disk of clypeus almost hairless, raised bristles only laterally and on
paraclypeal area. Antennal scape and pedicel reddish-brown, the former with thick yellow
mark anteriorly, segment 3 entirely black, except two minute anterolateral spots, one
at base, the second at apex; succeeding segments brown, their anterior faces ochraceous.
Long pubescence on genal area, lower parts of thorax and basal segments of legs, white.
Intero-ventral lamella at apex of gonocoxite placed transversely (Fig. 45), forming an
undulated plate, hollowed out within; when viewed from the inside, expanded basally,
projecting inward with strongly convex margin, distal portion tapering rapidly to a
blunt apex, the straight outer border of the lamella narrowly folded in almost for its
whole length. Gonostylus slightly surpassing lamella, at first slenderly lanceolate, then

M. A. LIEFTINCK: Review of Asiatic Habropoda 199

„lu
y di
|
il

17
/
Ji

=
SSS Aw ws

Nie
Ny N

4,
7
5
Ce
2
+
2
2
-
-
2
2
=
=

WH anas ANS

\u
I
I,

Fig. 42-45. Habropoda sutepensis CkIl., & structures, Thailand; Fig. 42, exterior view of left hind
tibia-basitarsus; Fig. 43, ventral view of seventh tergite; Fig. 44, seventh and eighth sternites,
exterior view; Fig. 45, left gonoforceps and penis valves, partial ventral view

broadening gradually, spatulate, the apex suddenly narrowing again and ending in a
blunt point (not shown in Fig. 45).

Female. — The brief diagnosis of this sex has been copied in the specific key. I am
unfortunately unable to give a more detailed description. The two topotypical specimens
which I had formerly registered — but failed to consider — in the Hymenoptera depart-
ment of the British Museum, could not be retraced when last inspecting the collections,
in October 1973.

This little species is known only from the type locality. Cockerell (1929) made the
interesting observation that the melectine Protomelissa habropodae (Ckll.), which it
greatly resembles, occurred together with many Habropoda sutepensis, stating that the
former is doubtless its parasite (see also Lieftinck, 1972 : 257, 281).

Habropoda hookeri Cockerell
Fig. 55

Habropoda hookeri Cockerell, 1920, Ann. Mag. Nat. Hist. (9) 6: 202—203 (2 Simla, 7000 ft.
Oct. 1907 (H.M.L.), Fletcher; and Mussoorie, 7000 ft., Aug. 1906, Fletcher). — Lieftinck, 1966,
Tijdschr. v. Ent. 109: 146 (notes).

Material examined. — North India, Uttar Pradesh: 1 9, with typewritten label

200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

“Simla 9.98”, and “P. Cameron Coll. 1914—110” (print), unidentified (BM).
Male unknown.

It is of interest to reproduce the original description of this bee, which runs as follows:

‘“@. — Length about 13 mm.
Robust, with abundant pale ochraceous-tinted pubescence, not mixed with black; clypeus
strongly rugose, very prominent, faintly keeled on apical half, with a transverse yellow
band, broadest in middle, just before the apex; first r.n. joining second s.m. before
the end.

This nearly agrees with H. montana Rad., as described by Bingham, but must certainly
be distinct, as the hind tibiae are not specially broadened or modified (their spurs are
ferruginous and very long), and their inner side, instead of being bare and smooth,
is densely covered with brownish hair. Other salient characters are: — Labrum with
ferruginous spot on each side of base; greater part of mandibles pale yellowish or cream-
colour; tegulae pale rufo-testaceous; hair on outer side of middle and hind tibiae
yellowish white (not bright ferruginous), some black hair near base of tibiae; hair
on inner side of hind basitarsi dark chocolate; wings brownish; nervures dark fuscous
(not testaceous); hind margins of abdominal segments broadly pallid; fifth segment
with a pale reddish fringe.

Simla, 7000 ft., Oct. 1907 (H.M.L.), Fletcher, 2 = type. Mussoorie, 7000 ft., Aug.
1906, also Fletcher, 2.

Dedicated to the memory of Sir Joseph Hooker.”

As pointed out earlier (Lieftinck, 1966), the characters distinguishing hookeri from
radoszkowskii, as given by Cockerell, apply to the leg structure of the males, and since
Cockerell had no male of hookeri, he was, of course, all abroad. Cockerell’s females,
including the type from Simla, could not be recovered in the British Museum (Nat.
Hist.) collection. However, the unexpected discovery of a single unidentified female
from Simla, mixed with various other Habropoda, enables me to give a better impression
of this bee. As it tallies the original description in almost every respect, the latter is
here copied verbatim.

The most important features of the present female are mentioned in the key. Chiefly
characterized by the almost white scopal hairs, the lustreless face, and the transverse
ivory mark bordering the clypeus anteriorly. Proboscis retracted, galea short, measuring
a little over 2 mm, not quite reaching base of fore coxae in repose. Long hairs covering
genal area, occipital region (mostly), the thorax ventrally, and the marginal fringes
of the abdominal sternites, almost white. Hair fringing posterior ridges of femora
likewise whitish, those at trochanter and femur of hind leg very short. Basitibial plate
brown, broadly rounded, surface flat with narrow upturned margin, its base covered with
dense yellowish tuft from apex of femur. Tibial scopa with short, linear stripe of
brownish hairs restricted to base of dorsal ridge, otherwise whitish. Penicillus of hind
basitarsus and most of the distitarsi ferruginous; claws tipped with dark brown. Wing
membrane lightly tinged with greyish yellow; 2nd submarginal cell almost square.

The “faint keel” on apical half of clypeus, mentioned by Cockerell, is unapparent
in the present specimen, there being only two short and low parallel ridges in the
median line, before the apex. Pygidial plate, Fig. 55.

The affinities of H. hookeri are obscure and must remain so until the male becomes
known.

M. A. LIEFTINCK: Review of Asiaiic Habropoda 201

Habropoda bucconis (Friese)
Fig. 46—49

Anthophora bucconis Friese, 1911, Verh. zool.-bot. Ges. Wien, 61: 127 (9 & Tainan, Formosa).
Strand, 1913, Suppl. Ent., 2: 51 (4 2 Formosa, addit. descr., partim!).

Type material examined. — Taiwan (Formosa): 1 4 (lectotype by present selec-
tion), “Tainan, Formosa/Anth. bucconis Fr., det. H. Friese 1910/Type” (print on dark
red), “Habr. bucconis, det. H. Hedicke 1942” (MNB).

Further material examined. — Taiwan (Formosa): 2 9, “Tainan, Formosa, Type”
(print on orange), det. H. Friese & H. Hedicke, evidently syntypes (MNB); 1 g'
(diss, Fig. 46-48), ‘’Taihorin, Formosa, H. Sauter 1911/7.xii, Anthophora bucconis
Fr., det. E. Strand/Dr. E. Strand 1913—249”, cpd. with lectotype, M. A. Lieftinck
(BM); 1 1 9, both labelled “Tainan, Formosa, Type” (print on orange), “Anth.
bucconis, ex coll. H. Friese”, evidently syntypes (SMF); 1 ¢ 2 9, “Hoozan, Formosa,
xii.1909 & i.1910, H. Sauter, A. bucconis, det. Friese & Hedicke” (MNB); 1 4, “Pilam,
Formosa, i.1908, H. Sauter’ (MNB); 1 9, “Polisha, x.1909, A. bucconis”’, det. Friese
(MNB); 1 2, “Formosa, Takao, xii.1908, Sauter” (print, ML).

When Strand (1913) writes: “Bei den meisten gg ist das ganze Untergesicht
gelb....”, he is certainly wrong, inasmuch as the genuine bwcconis was mixed up with
the similarly pubescent — but unrelated — fainanicola, a species that was also before
him at that time. This confusion is evident from his further remark: “Clypeus hat an
der Basis jederseits einen kleinen schwarzen Längsstrich”. In the male of bzcconis the
clypeus is brownish-black marked with a thick, yellow, T-shaped spot, the transverse
bar on top of the T being narrower than the stem, which is distinctly broadened ant-
eriorly. The females also can be distinguished by the clypeal marks (Strand: “Der helle
Clypeusfleck der 9 © kann fehlen”). True bucconis, however, has a more protuberant
face and the clypeus bears a triangular yellow median spot in front of the anterior border,
whereas in fainanicola the clypeus is invariably unicoloured black, or dark brown.

Though very similar superficially, these two Taiwanese species are altogether different
structurally, belonging to two different species groups.

For a description of this bee, see the keys to both sexes and the comparative notes
as contained in the diagnosis of its nearest relative, zmitatrix, in the next pages. Indeed,
were it not for the different structure and pubescence of the hind legs, and the slightly
dissimilar form of the hidden sternites and genitalia of the male, the evidence for
considering these two taxa specifically rather than subspecifically distinct, would seem
to be less strong than, for example, in the case of bucconis and the previously described
tainanicola. Unfortunately, these features are unisexual, the recognition of females being
more difficult, necessitating a careful comparison of all characters.

As far as our locality records show, bucconis and fainanicola are probably completely
allopatric in Taiwan, as suggested above, at the end of the discussion of the latter.
Another fact separating them, is the difference in flight dates, bucconis being principally
a late season bee, collected from October till January, whereas tainanicola, as we have
seen, is an early spring species. In respect of seasonal occurrence, bucconis thus appears
to be neatly affiliated with its ally of the Chinese continent, /mitatrix, of which all
specimens were collected in the autumn, between August and November.

Note. — Dr. Maa tells me (in litt.) that all old Taiwanese (Formosan) localities
as found in the literature are situated in low hilly country or near the coast; the names

202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

TRENT

Fig. 46—49. Habropoda bucconis (Friese), structures, Taiwan; Fig. 46, exterior view of & left hind

tibia-basitarsus; Fig. 47, eighth and seventh sternites of 4, exterior view; Fig. 48, left gonoforceps

and penis valves, partial ventral view, with detached right gonostylus, dorsal view; Fig. 49, pygidial
plate of 2, dorsal view

of several of these places have since been changed, as follows. Hoozan = Fengshan
(low hills); Pilam = Taitung (sea level); Polisha = Puli (450 m, but up to 1000 m

alt. further inland); Tainan = unchanged (low country); Taihoku

= Taipei (low
country); Taihorin = Tailin (low country); Takao = Kaohsiung (low hills). Owing
to serious deforestation and other drastic changes through human agency, the natural

surroundings of south Taiwan are nowadays very different compared to Sauter’s time.

M. A. LIEFTINCK: Review of Asiatic Habropoda 203

Habropoda imitatrix spec. nov.
Fig. 50—54

Material examined. — SE China, Fukien prov.: 78 g' (2 diss.), 75 ©, Shaowu
distr., Tachulan, 1000—1200 m, ix-x.1942—43 (0 diss, 8.ix.1943, Fig. 50—53);
4g 2 Q, Shaowu distr., Outou, 16.x.1943; 2 g 1 9, Shaowu distr., Lichiatun &
Tengchiatun, 16—18.x.1943; 2 & 7 9, Chungan distr., Sanchiang, 1000 m, 17.x.1941
& 24—31.x.1942; 1 9, Chungan distr., Kuatun, 1200—1500 m, 28.x.1942; 8g 1 9,
Kienyang distr., Hwangkeng, 500—800 m, 18.x.1941, 4.xi.1942 and 15—17.x.1943;
all T. C. Maa. SE China, Kiangsi prov.: 1 g 2 9, Kwantseh, Kwanmentshih, viii. 1943,
T. C. Maa. Holotype g' and first described 9 : Shaowu, Tachulan, 8.iv.1943 (9° diss.),
and same loc., 1—15.x.1943 (2) (ML); paratypes in various other collections.

A homogeneous series of both sexes, which show little or no variation, except in size.
Males average slightly smaller than females, but exhibit the same uniform colour design.

Stature of H. bucconis and radoszkowskii, and only superficially resembling tainanicola
cum subsp., described before.

Male. — Proboscis of moderate length, galea reaching back in repose to as far as
middle of fore coxae. Labrum dark brown, with lighter brown basal tubercles; only a
trifle broader than long, disk shining, irregularly, rugosely punctate, anterior border
somewhat raised and swollen on either side of a tiny, shallowly crescentic emargination,
the edges of which are prominent, almost rectangulate; pile long, silky silvery white,
all hairs directed forward, as are the long, stiff marginal bristles emerging from the
emargination. Mandibles ivory, a brown line at base as well as the apices, black. Malar
area light brown, a little longer than pedicel of antenna and almost 114 times as long
as in bucconis. Antenna dark brown, apex of segment 3 with orange spot and 3—13
ochraceous anteriorly; scape broadly striped with yellow in front, scarcely longer than 3.
Ocelli placed in equilateral triangle, the laterals more distant than in bucconis. Vertex
and mesonotum rather shining, surface of the former finely chagreened, impunctate,
centre of mesonotum sparsely, superficially punctate, the sides much more closely so.
Tegulae light brown, shining, basal half clothed with longish decumbent hairs, directed
toward centre and not quite concealing surface. Clypeus dark reddish brown, the cream-
coloured median bar halter- or I-shaped, much narrower than in bucconis, transverse
basal portion of this spot occasionally reduced and tapering basad to a mere stripe; long,
silvery decumbent pile at sides converging, directed obliquely cephalad, the tips of hairs
meeting in the median line and hiding most of the surface, sides moreover with sparsely
distributed, long erect hairs, longest at side-angles; yellow paraclypeal areas clothed
densely with fine, pale pubescence. Occiput and genal area white-haired, the former in
addition fringed with much longer, raised dark hairs. Legs dark brown, hind basitarsus
darkest, mediotarsi lighter brown, last segment ferruginous; pubescence predominantly
pale brown, long, raised and almost white at all femora, the outer fringes of fore and
mid tibiae also white, but long hairs fringing lower face of hind tibiae darker; fringes
at dorsal ridge of hind tibia light orange, increasing in length from base to apex, the
distal hairs much longer than in bzcconis and tipped with black. Spurs pale. Hind
basitarsus almost parallel-sided, surface shining, sulcate, scatteredly punctate (similar to
bucconis); fringe at dorsal crest longer and denser than in that species, brownish black
or black instead of short and pale-coloured. Wing membrane yellowish, venation as
variable as in bucconis and radoszkowskii; 2nd submarginal cell frequently almost square,

204

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

My yy
1

Il
dl

SA
Raz,

PE rn >>

HALF

> AAA

Fig. 50—54. Habropoda imitatrix spec. nov., structures, Fukien; Fig. 50, exterior view of & left
hind tibia-basitarsus; Fig. 51, ventral view of & seventh tergite; Fig. 52, eighth and seventh sternites

of &, exterior view; Fig. 53, left gonoforceps and penis valves of &, partial ventral view, with
detached right gonostylus, dorsal view; Fig. 54, pygidial plate of ©, dorsal view

receiving recurrent nervure at, or (more often) a little before, its distal side. Punctation,

colour and pubescence of abdominal tergites and sternites as in bucconis, hind margins
of sternites 2—5 straight, the conspicuous, long and dense, apical hairs pure white,
similar in the two species. Sternites 7 and 8 as in Fig. 52, the microsetae covering ventral
face of 7 strongly curved, rather similar in arrangement to bwcconis. Intero-ventral
process at apex ot gonocoxite, in full ventral view, broader and more plate-shaped than

M. A. LIEFTINCK: Review of Asiatic Habropoda 205

shown in Fig. 53, a little hollowed out dorsally, subrectangular in outline, but widest
and broadly rounded mesially at base, the apex folded ín and carrying a strong recurved
ridge, which does not reach the base of plate; whole ventral surface covered with
microsetae. Gonostylus subequal in length to plate (Fig. 53).

Female. — Labrum similar to bucconis: distinctly broader than long (hence shorter
than in male); apex of the slightly projecting anterior border divided transversely into
a thin lower plate and the proper margin immediately above it, the latter being distinctly
swollen and upturned on each side of the middle, forming a pair of black tubercles,
the median tuft of bristly hair emerging from the sulcus between the two plates. Clypeus
and supraclypeal area invariably darker brown than in bzcconis, the ivory spot, though
varying in size, less sharply defined than in that species. Antennae also darker, and of
uniform colour. Body punctation and vestiture, as well as shape and pubescence of legs,
practically identical in the two species; basitibial plate also shaped similarly, almost
circular in outline, disk quite flat, clothed evenly with fine, appressed golden tomentum,
border slightly thickened, almost black. Pygidial plate light brown, darker towards apex,
surface only very slightly convex, rather dull, finely transversely wrinkled, shaped much
as in bucconis, which has the apical portion of the plate rather more elevated, showing
markedly depressed margins (cf. Fig. 49 and 54).

In my key to the identification of the males, the couplets leading to bzcconis and
imitatrix run out far apart, a diversion entirely due to the intentional selection of only
two characters, viz. leg structure and vestiture of the abdominal sternites. In view of
other characters separating these two species, it is difficult to judge whether too much
stress has, perhaps, been laid on the above features. The male terminalia are very much
alike. It should be noted that the shape of the appendages of the gonocoxite, when
viewed from below (Fig. 53), probably depends largely on the amount of folding, or
even shrinkage, of these delicate setiferous lamellae. It follows that their form can hardly
be used as a means of distinguishing between these two species.

Habropoda radoszkowskii (Dalla Torre)
Fig. 56—62

Podalirius radoszkowskii Dalla Torre, 1896, Cat. Hym., 10: 285 (nom. nov., nec Anthophora
montana Cresson, 1869, Trans. Amer. Ent. Soc. 2: 290, ® ). — Friese, 1897, Bienen Europa's, 3:
305—306 (2 & latin diagnosis, sec. Radoszk.). — Bingham, 1897, Fauna Brit. India, Hym. 1: 522
(2 only!), fig. 177 (2 insect), “The Himalayas from Sikhim to Simla, above 4000 ft.”

Habropoda montana Radoszkowski, 1882, Wiadom. nauk przyrodz. Warszowa, 2: 77 (2 6 Hima-
laja).

Habropoda radoszkowskii: Lieftinck, 1966, Tijdschr. v. Ent., 109: 144—145 (notes or: allied
species).

Type material examined. — North India: 1 4 (lectotype by present selection),
1 9, labelled “HIMAL” (print), “coll. Radosz.” (print), both with “montana” (in
Radoszkowski’s hand), “Typus” (print on light red), “Habropoda montana, det. H.
Hedicke” (MNB). 1 & (syntype montana Rad.), “Cotype” (print on yellow disk),
“HIMAL” (print on yellow), ‘“Habropoda montana Rad. 4” (Radoszkowski’s writing),
small gold disk (BM).

Further material examined. — NE India: 14 2 9, “Sikhim, Coll. Bingham,
montana, det. H. Hedicke” (MNB); 4 9, “Assam, Khasia Hills 96—135’ (BM);
1 2, “Assam, Shillong, 10.03, Coll. Bingham, H. radoszkowskii 9” (unknown hand),

206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

“montana, det. H. Hedicke” (MNB); 1 9, “India, Darjeeling, 6.xi.1920, leg. Brunetti,
BM 1927—184” (BM); 1 9, “Sikkim, Darjeeling, 7000 ft., 4.94” (Bingham’s label),
“H. radoszkowskii D.T., det. Bingham” (BM); 1 9, “Sikhim, 4.94, H. radoszkowskii
Dall. Torr.” (Bingham’s hand) (OUM); 1 9, “Sikkim, Darjeeling, 7000 ft., 4.94,
Bingham Coll. “(OUM); 1 g', “Sikkim, 4/95 31/5, Coll. C. T. Bingham 96—30”
(BM); 5 9, “Darjeeling, 6—7000 ft, 10.93 & 4.5.94” (BM). North India:
2 d, “Kumaon, U.P., 10.89, Miss A. Brook, Habr. radosz. Coll. Bingham” (Bingham’s
hand) (BM); 2 2, “Kumaon, U.P., W. Almora Div., Nov. 1919, H.G.C. “(BM);
2 2,” Punjab, Dalhousie, Oct. 1917, N.B. Kinnear’ (BM). East Nepal: 13 g
(2 diss.), 10 9, Taplejung [Taplang Jung] Distr., deep river gorge, 5200’, and mixed
shrubs in deep gorge, 5700’, between Sangu & Tamrang, x-x1.1961 (2 g'), Sangu,
6200’, mixed vegetation by stream in gully, ix-x-1961, and yellow blooms of cultivated
Composite, 16-29.x.1961, edge of mixed forest above Sangu, 6500’, 17.x—1.xi.1961
(1 & diss.); Sangu, 6200’, blooms of wild cherry, 15—18.xi.1961 (4 & 3 2); old
mixed forest above Sangu, 6200’, 25—28.x.1961 (1 & 2 2); Sangu, 6200’, on minute
flowers of wild shrub, 25.xi.1961, clearing in small wood (2 g' 5 2); all specimens
R. L. Coe leg., Brit. Mus. East Nepal Exp. 1961—62 (BM; 2 g' 2 9 ML).

Owing to some discrepancies in the existing descriptions, the confusion around this
species has been considerable. In my 1966 paper, I have commented already upon some
of these problems, which could mostly be solved since then. One of Bingham’s males,
described by him from Simla, is certainly not radoszkowskzi, but its identity must remain
uncertain.

Though several features obviously relate this species fairly closely with bucconis and
imitatrix, both sexes of this bee are further distinguished by the shorter thoracic and
abdominal pubescence, so that more of the dark integument is exposed, especially at
base of the first three or four abdominal tergites. Fresh males in our series are as vividly
coloured as those of the other two species, whereas the pile covering the thorax of the
females has a more sallow, greyish orange, tint.

Male. — Proboscis relatively short, galea in repose not quite reaching fore coxae,
measuring only 2.8 mm. As compared with the preceding species, the anterior border of
the labrum is a little more convex on either side of the crescentic emargination; oblique
ante-apical ridges deep black, more pronounced and meeting in the centre of the emar-
gination; disk shining, with few superficial punctures, though densely hairy. Face
shaped and coloured similarly to zmitatrix. Yellow clypeal mark intermediate in shape
between that of bucconis and imitatrix: the constricted part as broad as the yellow mark
on the antennal scape. Antenna dark brown, but all segments lighter anteriorly: scape
citron-yellow, the distal two-fifths of 3 and most of 4 conspicuously orange, the next
segments gradually becoming less bright, rather more ochraceous, in front. Legs more
robust and all tibiae more swollen than in the preceding species. Spurs brownish yellow.
Sole of hind basitarsus exactly four times as long as its greatest breadth, apex markedly
swollen and, when viewed obliquely from the inside, hollowed out internally (Fig. 57).
Wing venation variable, but 2nd submarginal cell relatively longer than in the allied
species and recurrent nervure never quite interstitial. Vestiture of gastral sternites differ-
ing from that seen in bucconis and imitatrix, approaching tainanicola more closely in
this respect. The tufts of soft raised hairs at the sides are of great length, silvery white
and most conspicuous at 3; from then on rearward, the marginal hairs gradually become
shorter and wavy toward the median area of sternites, acquiring a yellowish tint, those

M. A. LIEFTINCK: Review of Asiatic Habropoda

Mehdi

À
S
Ÿ
=
=

Fig. 55. Habropoda hookeri Ckll., Simla, apical portion of © pygidial plate, dorsal view, greatly

enlarged. — Fig. 56—62. H. radoszkowskii (D.T.), structures, Nepal; Fig. 56, exterior view of &

left hind tibia-basitarsus; Fig. 57, ventral aspect of basitarsal “sole” of same (slightly more

enlarged); Fig. 58, ventral view of 4 seventh tergite; Fig. 59—60, seventh (59) and eighth (60)

sternites of 4, exterior view; Fig. 61, left gonoforceps and penis valves of 4, partial ventral view;
Fig. 62, pygidial plate of 9, dorsal view

207

208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

covering tne apical portion of 5 being shortest, forming a transverse brushy belt of very
dense, light golden pubescence, flanked by the equally dense (but much longer) white
tufts at the sides of the sternite. Sternite 6 shaped similarly to bucconis and imitatrix,
the bluntly triangular tip clothed densely all around margin with minute erect hairs.
Sternites 7 and 8, Fig. 59—60. Intero-ventral process at apex of gonocoxite more
elongate (cf. Figs. 53 and 61), its rounded basal portion longer, projecting cephalad,
the whole plate concave dorsally; viewed from below, a narrow ridge extends from the
margin inward (not shown in Fig. 61). Gonostylus about equal in length to the plate,
apex broad, abruptly expanded and fringed with long, radiating bristles.

Female. — Antenna dark brown, as in male, but the orange apical spot in front of
segment 3 is reduced in size, the flagellar segments (except 4) being less brightly
ochraceous than in the other sex. Basitibial plate shaped and coloured as in bucconis and
imitatrix, with the same compact quiff of downbent golden hair at apex of femur
overlying its base.

Not all Indian localities mentioned by Radoszkowski and Bingham are precisely
known, but the species seems to be widely distributed in the Himalayan region. In Nepal
H. radoszkowskii was found in occupation of the same territory as several of its allies,
two new species, viz. apostasia and pelmata, having been collected along with it in the
Taplejung District, near Sangu, and all at one time (September till November) of the
same year. The nearest ally of radoszkowskii would seem to be H. plantifera spec. nov.,
from Assam, described hereafter.

Habropoda plantifera spec. nov.
Fig. 63—67

Material examined. — NE India: 1 4 (holotype, diss.), Assam, labelled “Shil-
long, 10.03” (written), “Turner coll. 1912—111” (print), in drawer sub radoszkowsku
(BM).

In the specific key to the males, couplet 7 carries this species to group 10', which
comprises radoszkowskii and two of its immediate allies, viz. plantifera and pelmata,
all characterized by the remarkable sole-shaped thickening on the hind basitarsus. There
is only a single male.

As compared with radoszkowskii, the following additional peculiarities can be
recorded.

Male (holotype). — Proboscis extended, but glossa retracted; galea equal in length
proportionally to that of radoszkowskii, measuring 3.0 mm. Labrum subrectangular, a
little broader, the anterior border nearly straight, hardly thickened, median incision
unapparent, replaced by a pair of minute dark teeth; disk almost flat, smooth and
shining, with few superficial punctures at base only; colour light brown, long hairs and
apical fringe golden yellow. Clypeus as described; facial pubescence all pale, except
long raised paraclypeal, interocellar, and occipital bristles, which are brown. Antenna
dark brown, apical spot on anterior face of 3 and all along remaining flagellar segments
somewhat lighter brown. Long pubescence covering genal area, basal segments of legs,
and ventral parts of thorax and abdomen, palest yellow, not pure white. Sole of hind
basitarsus over four times as long as its greatest breadth (100 : 24.2), apex a little less
swollen and, when viewed from the inside, correspondingly less hollowed out, than in
radoszkowskii. Wing venation lighter brown, the membrane of fore wing more deeply

M. A. LIEFTINCK: Review of Asiatic Habropoda 209

ex
\)
EN SS
\

fi
7
AA
7
Je
z
2
7
7
7

Ä
RMS

WOS

ON
DW

TS
In

Fig. 63—67. Habropoda plantifera spec. nov., structures, & holotype Shillong; Fig. 63, exterior

view of left hind tibia-basitarsus; Fig. 64, ventral view of seventh tergite; Fig. 65—66, seventh (65)

and eighth (66) sternites, exterior view; Fig. 67, left gonoforceps and penis valves, partial ventral
view

stained with yellow, especially toward apex; costal side of 2nd submarginal cell markedly
shorter than anal side; recurrent nervure entering cell well before distal side of the
latter. Vestiture of thorax and abdominal tergites a little longer, the tergites distinctly
more evenly and densely clothed with warm orange pubescence; arrangement of sternal
hairs much as in radoszkowskii. Apex of tergite 7 slightly longer and narrower than
in that species. Sternites 7 and 8, Fig. 65—66. Intero-ventral lamella at apex of
gonocoxite very different in shape, the broadened subbasal portion not projecting
cephalad, but inward, and more rounded off, the apex after a constriction being broadly

210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

rounded and distinctly ridged interiorly (not shown in Fig. 67). Gonostylus equal in
length to lamella, slender and strongly incurved basally, its apical portion foot-shaped,
the “heel” fringed with long hair, as in the figure.

The name of this species refers, of course, to the form of the hind basitarsal plate.

Habropoda pelmata spec. nov.
Fig. 68—71
Material examined. — Nepal: 1 & (holotype, diss.), E Nepal, Taplejung
[Taplang Jong} Distr., Sangu, ca. 6200’, clearing in small wood, 25.xi.1961, R. L. Coe
coll, Brit. Mus. East Nepal Exp. 1961—62 (BM).

This is the second nearest ally of H. radoszkowskii, quite distinct, but unfortunately
represented only by a single male not in a too good condition. It has lost most of the
antennae, the wing margins are torn, and the discoloured pubescence has been partly
rubbed off. The diagnosis in the key can be extended as follows:

Fig. 68—71. Habropoda pelmata spec. nov., structures, 4 holotype Nepal; Fig. 68, exterior view
of left hind tibia-basitarsus; Fig. 69, ventral view of seventh tergite; Fig. 70, eighth and seventh
sternites, exterior view; Fig. 71, left gonoforceps and penis valves, partial ventral view

M. A. LIEFTINCK: Review of Asiatic Habropoda 211

Male (holotype). — Proboscis retracted, galea equal in length to that of plantifera,
reaching to base of fore coxae in repose. Labrum also similar, brownish-yellow, becoming
darker apically; no median incision, but ante-apical teeth less approximated; long pile
covering disk and sides of paraclypeal area white. Basal two-thirds of clypeus rather
dull, microscopically tessellate, quite similar to plantifera, which also has the apical
portion indefinitely marked off, only its surface being more shining and the low rugae
less distinct, than in plantifera. Antenna dark reddish-brown, segment 3 in front with
vestigial basal and a somewhat larger apical ferruginous spot, (rest missing). Punctation
in the centre of mesoscutum much less dense than at sides, though even here the
punctures are a little smaller than the finely tessellated spaces separating them (apparent-
ly quite similar to plantifera, where much of the ground is hidden by pubescence). Body
hair sallow, obviously rather discoloured, pale orange-yellow, hence much less bright
than in the preceding species. Legs distinctly more slender (cf. Fig. 63 and 68), the
tibiae and basitarsi with shorter hair. Sole of hind basitarsus noticeably narrower than in
plantifera, though well marked off, 5.8 times as long as its greatest breadth (100 :
17.24), cylindrical with tapered apex; in profile, the presence of this inferior plate is
revealed by a slight concavity of the lower margin, as well as by a gradual broadening
toward the end of the segment itself. Wing membrane pale yellow; costal side of 2nd
submarginal cell only little shorter than anal, the recurrent nervure is received slightly
before the end of the cell. Abdominal pubescence long and thin, partly rubbed off
so as to expose the broad, contrastingly pale-coloured posterior borders of all tergites;
sternal pubescence as in plantifera. Tergite 7 and hidden sternal plates 7 and 8, Fig. 70.
Intero-ventral lamella at apex of gonocoxite shaped much as in the preceding species,
but in full ventral view the plate is on the whole broader, less deeply concave inward
about halfway length. Apex of gonostylus very similar in outline, though less distinctly
foot-shaped, than in plantifera (Fig. 71).

Female unknown, unless the ambiguous “apatelia” from Nepal may turn out to be
the same species?

The males of the next two species, deiopea and apatelia, bear a close prima facie
resemblance to pelmata, described above and are undoubtedly also members of the
radoszkowskii group. As we have seen, they all share the curious smooth and hairless
expansion on the underside of the hind basitarsus, which here forms a convex, lanceolate,
or rather torpedo-shaped plate, a modification displayed in a more exaggerated way
by the males of radoszkowskit, plantifera and pelmata, in which the plate is definitely
sole-shaped.

The two species in question can be easily recognized from the other members of the
group by lacking a yellow spot in front of the antennal scape. They are much alike,
and if no fair series had been available, recognition would have been difficult, because
little is still known of the individual variation of each. As it is, the most striking
features exhibited by a total of 21 males were carefully compared, and this has resulted
in the segregation of at least two closely similar species, discussed hereafter. In regard
to the females, we have only a single individual of either species, and these could be
matched with their males with reasonable certainty.

212 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Habropoda deiopea (Cameron)
Fig. 72—77

Anthophora deiopea Cameron, 1897, Mem. & Proc. Manchester Lit. & Philos. Soc. 41 (4): 127—
128 (2 4 Mussouri, Rothney).

Type material examined. — NW India, Uttar Pradesh: 1 g' (diss, Fig. 75),
“Mussoori”, with small square label “m”, and “Osmia deiopia {sic} Cam. Type”, in
P. Cameron’s hand, and “Habropoda montana Radosk, J”, in Bingham’s writing.
Lectotype A. deiopea Cam., by present designation (ex Rothney coll, OUM).

Further material examined. — North India, Uttar Pradesh: 9 g' (one diss.,
Fig. 73—74), each with typewritten labels “Simla 9.98”, and “Coll. G. Nurse Col-
lection 1920—72” (print), all unidentified (BM). 1 9, “India, Uttar Pradesh, Mus-
soorie, Lal Tibba, Alt. 7500’, 19.x.1964, S. W.T. Batra, on Salvia glutinosa L., Habro-
poda radoszkowskyi D.T.’’, det. Batra (UKL)

The series of males from Simla in the British Museum, were standing side by side
with a single female in one drawer and bear identical typewritten locality labels, but
originate from different collections. These males are not conspecific with the female,
which corresponds closely with the original description of H. hookerz Ckll.; see that
species. The males are exactly similar to Cameron’s male of A. deiopea, marked “Type”.
As the original female deiopea is nowhere to be found and has apparently been lost,
the male is here designated lectotype. The head markings, colour of pubescence, as well
as the leg structure, are all precisely as in a specimen from Simla, the same congruity
being manifest on comparing the internal sternal plates and genital organs, which were
dissected out in both. These structures are here shown for one of the Simla males in the
British Museum (Fig. 72—74).

Male. — Though resembling H. apatelia fairly closely in the uniformly coloured body
pubescence, deiopea can be recognized, apart from the distinctives enumerated in the
key, by its broader forms and, on dissection of the terminalia, by the shape of the
gonostylus, the apex of which is emarginate (Fig. 76); the remaining differences are
slight and difficult to paraphrase. In the lectotype the 7th tergite is shaped similarly
to that of a Nepalese specimen of apatelia, shown in Fig. 79. It should be noted that,
owing to a slight distortion of the genital capsule, the intero-ventral lamella of the
gonocoxite, shown in situ (Fig. 75) for the lectotype, had to be drawn in a somewhat
twisted vertical position; when viewed from the external side, it is shaped exactly as in
Fig. 76, taken from a Simla male.

Female. — The present specimen from Mussoorie is topotypical and fits the de-
scription of the female in every respect. Here follow Cameron’s diagnoses of the two
sexes (loc. cit.: 127—128).

“Nigra, longe dense pallide hirta; capite nigro g' facie alba.

Long. 13 mm. 9.

Hab. Mussouri (Rothney).

Head black, thickly covered with long pale grey hairs, especially on the front and
vertex; the labrum fringed with golden hairs; the mandibles ferruginous, black at the
apex. The vertex behind the front ocellus bare, shining, broadly depressed. Thorax
thickly covered with long grey hairs all over. Legs: the femora and tibiae dark rufous;
the former sparsely covered with long white hairs, the front four tibiae covered densely
behind with pale fulvous hairs; the hairs on the hinder tibiae much longer, thicker,

M. A. LIEFTINCK: Review of Asiatic Habropoda 213

Fig. 72—77. Habropoda deiopea (Cam.), structures, Simla; Fig. 72, exterior view of & left hind

tibia-basitarsus; Fig. 73—74, seventh (73) and eighth (74) sternites of &, exterior view; Fig. 75,

genital capsule of & lectotype (Mussoorie), ventral view; Fig. 76, exterior view of right intero-

ventral lamella and gonostylus of 3 gonocoxite, more enlarged (Simla); Fig. 72, pygidial plate of
9, dorsal view (Mussouri)

and of a brighter fulvous tint; the tarsi rufous, thickly covered with long golden hairs at
the base. Wings hyaline, with a faint fuscous tinge; the costa and nervures blackish; the
first recurrent nervure is received shortly before the second transverse cubital nervure;
the second is interstitial. Abdomen above and at the sides thickly covered with long pale
fulvous hairs; the penultimate segment rufous at the apex; the apical ferruginous, black
at the apex, the base closely transversely striated; the sides, especially towards the apex,

214 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

broadly furrowed; abdominal segments black, the base and apex broadly ferruginous;
the segments at the apices thickly fringed with fulvous hairs.

The 8 is covered all over with long hoary hairs; the clypeus, except at the sides and
the inner orbits, cream-yellow; the extreme apex piceous, the mandibles cream coloured;
the tips black, ferruginous in front of the black; the labrum black, covered with white

hairs; the ventral segments are coloured as in the © ; this being also the case with the
legs, which bear long white hairs.”

In general appearance, colour and size, the female from Mussoorie makes an excellent

match for our series of males, hence I do not doubt to have associated the sexes correctly.
Pygidial plate, Fig. 77.

—-
SANARE SEN eases

{

N
N Sa
ERS

\ 2
N

N =
N È
N

Fig. 78—84. Habropoda apatelia spec. nov., structures, Darjeeling; Fig. 78, exterior view of & left

hind tibia-basitarsus; Fig. 79, ventral view of 3 seventh tergite; Fig. 80, eighth and seventh sternites

of 4, exterior view; Fig. 81—83, left gonoforceps and penis valves of 4, partial ventral view (81),

interior (82) and ventral (83) view of right gonocoxal appendages of same, more enlarged; Fig. 84,
pygidial plate of 2, dorsal view

M. A. LIEFTINCK: Review of Asiatic Habropoda 215

Habropoda apatelia spec. nov.
Fig. 78—84 and 85—89

Material examined. — NE India: 7 & (one diss, holotype), 1 9, Sikkim, all
with identical printed labels “India Darjeeling 1920, Brunetti, B.M. 1927—184”, and
written on reverse side ‘“10.X” (BM, sub rowlandi), no identification labels. Holotype
d', unique ® and 4 Z' paratypes (BM), 2 g' paratypes (ML).

Further material examined. — Nepal: 4 & (one diss.), E Nepal, Taplejung
[Taplang Jong} Distr., old mixed forest above Sangu, ca. 6200’, 25—28.x.1961, R. L.
Coe coll, Brit. Mus. East Nepal Exp. 19961—62 (4 diss., BM); same area, Sangu,
ca. 6200’, clearing in small wood, 25.xi.1961, same coll. (2 g', BM); same area, above
Sangu, ca. ca. 6500’, evergreen scrub, 5—13.x.1961, same coll. (1 &, ML).

The following topotypical females are of uncertain identity and are only tentatively
placed here; they may be conspecific either with apatelia or pelmata, also described from
Nepal. Nepal: 11 9, E. Nepal, Taplejung [Taplang Jong} Distr., Sangu, ca. 6200’,
on minute florets of shrub, clearing in small wood, 25.xi.1961; and same area, old mixed
forest above Sangu, ca. 6200’, 25—28.x.1961, same coll. (BM, ML).

Male. — It should be emphasized that the characters given in the descriptive key

\
\

Fig. 85—89. Habropoda apatelia spec. nov, & structures, Nepal; Fig. 85, exterior view of left hind
tibia-basitarsus; Fig. 86, ventral view of seventh tergite; Fig. 87, eighth and seventh sternites, exterior
view; Fig. 88—89, left gonoforceps and penis valves, partial ventral view, with detached right
gonostylus, dorsal view (88), and interior view of right gonocoxal appendages, more enlarged (89)

216 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

were taken primarily from the Darjeeling males (including the type), and that only
the latter can be considered paratypes of apatelia. The remainder, from Sangu in Nepal,
differ so perceptibly from the typical series that they were first segregated from the
others and given a new name. However, considering that all but one are worn and
discoloured individuals, it was decided to keep them together. With better material and
the discovery of females, these Nepalese males may well prove specifically distinct after
all. The differences are briefly summarized in the key and structural details are given
niro 7080?

Female (Darjeeling). — Closely resembling deiopea, but differing from that species
in a number of details enumerated in the key.

Female (Nepal). — Of doubtful identity. Distinguished from the only authentic
female of apatelia in that the integument of all parts of the body is of a darker brown.
Clypeus with well defined ivory spot, usually forming an isosceles triangle, upon middle
of distal half just before anterior border, this spot occasionally narrowly framed in
reddish brown. IOD : OOD = 50 :100 approx. Length ratios of antennal segments
3, 4 and 5 = 100 : 30. 40. Pygidial plate shaped and sculptured much as in bucconis
(Fig. 49), but surface dull and apparently not striated.

Except for their smaller size, these females are inseparable from those of undisputed
radoszkowskii, with which they were collected simultaneously. As hinted at above, the
possibility remains that these bees can be associated with our new Nepalese Habropoda
pelmata, which also is a smaller-sized species.

Habropoda krishna Bingham

Habropoda krishna Bingham, 1909, Rec. Ind. Mus., 2; 366—367 (& Sikkim, Darjeeling, 7,000
ft.). — Cockerell, 1911, Proc. U.S. Nat. Mus., 39: 643 (note); Cockerell, 1920, Ann. Mag. Nat.
Hist, (9) 6: 202 (Khasia Hills, Sladen). — Lieftinck, 1966, Tijdschr. v. Ent., 109: 145 (note on
characters).

An enigmatic species. Type (ex Zoological Survey of India, Calcutta) probably lost.
Female unknown.

The original description is based on a single male, sent to the describer by Dr. Nelson
Annandale. It runs as follows:

“g. Black. Head, thorax, and abdomen densely covered with long, pale brownish
yellow pubescence turning to ferruginous golden on the posterior segments of the
abdomen, and mixed throughout with a sprinkling of black hairs, the surface beneath
more or less closely and finely punctured. Head: mandibles white; apical third rich
castaneous; base black; clypeus white; apical margin narrowly castaneous; sutures at the
sides and above narrowly black. Thorax: the wings hyaline; nervures and tegulae testa-
ceous. Legs: apical joints of the tarsi rufo-testaceous; claws black. Head as wide as the
thorax; the vertex broad, depressed; antennae comparatively long, second joint of the
flagellum attenuate at base, clavate at apex, as long as the succeeding three joints; ocelli
in equilateral triangle, the distance between the anterior and either of the posterior ocelli
greater than either of the posterior ocelli and the eye next to it; eyes large, prominent,
their inner orbits parallel. Thorax globose; mesonotum convex, scutellum and post-
scutellum slightly raised, median segment short, very convex above.

In some specimens not so densely pubescent as others, the pubescence forms distinct

M. A. LIEFTINCK: Review of Asiatic Habropoda 217

transverse bands on the apical margins of the segments beneath which the surface is
rufo-testaceous. Female unknown.

Length, — G', 12—13; exp. 24—27 mm.

Hab. — Sikhim: Darjeeling, 7, 000 ft.

Nearest to H. radoszkowskii, but differs in sculpture and in colour of pubescence.”

As follows from the above description, the relative lengths of the flagellar segments
of antennae, in combination with the distance ratios of the ocelli, are characters not in
accordance with any of the described species and at variance with all that is known about
the morphology of male Habropoda. Cockerell (1911) evidently knew this species only
from the description as he gave no further characters. In 1920, the same author claims
to have seen a specimen from the Khasia Hills bearing a manuscript name by Friese,
who dedicated the species to Mr. Sladen — an irrelevant observation. This example is
not in the British Museum collection.

The aberrant features mentioned above may be due to incorrect observation, because
everything else points to near relationship with H. radoszkowskii and its allies.

Habrophorula gen. nov.
Fig. 90—100

Of the Anthophorini. Medium-sized bee of elongate form, stature recalling Elaphro-
poda Lieft., but legs entirely unmodified, thin and slender, especially in male.

Head much broader than long in dorsal view; eyes moderate, widely distant, inner
orbits subparallel, slightly convex inward and diverging anteriorly (Fig. 90—91). Pro-
boscis long; last two segments of labial palpi short, fused together and ribbon-like;
maxillary palpus 6-segmented, the second segment longest. Labrum broader than long,
widest about middle, somewhat protuberant, anterior border not excised, its surface
smooth, lacking subapical ridges or tubercles (g'); or longer, almost square, though
otherwise similar ( 9). Mandible with single, subapical, interior tooth. Malar area
distinct, subequal in length to pedicel of antenna. Clypeus little protuberant, greatest
anterior depth in side view nearly equal to diameter of eye (g'), or even less (9).
Frontal carina low and very short (4 }, or a little longer ( 9). Vertex slightly raised,
surface convex, ocelli close together, forming an isosceles triangle. IOD:OOD =
49 :100. Antenna slender, third segment exceptionally short in both sexes, remaining
flagellar segments cylindrical, longer than thick (6th segment about 10 : 6).

Thorax short and small, globose. Le gs of simple structure, those of male long
and very slender; coxae unarmed, those of fore legs large, subtriangular, of mid leg
elongate-oval, of hind leg more or less triquetral in cross-section; trochanter of fore and
mid legs longer than broad, of hind leg squarish; all tarsal claws deeply cleft, the rami
subequal in length (3), or inner ramus shorter on fore, mid and hind tarsi: about
four-fifths, three-fourths and two-thirds as long as outer, respectively (2); arolia well
developed. Basitibial plate of female forming an equilateral triangle with rounded angles,
of male also well defined, subcordate with slightly produced, rounded apex. Scopa
of female rather short and thin, hairs plumose.

Wings short. Marginal cell of fore wing considerably longer than distance from its
apex to wing tip (100 : 62.5), free part of marginal cell rather shorter than rest of cell;
first s.m. almost twice as long as second but shorter than second and third united,
measured on anal side; second 5.72. squarish, a little higher than long, as in many

218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Habropoda, with costal side shorter than anal; third s.m. much longer, about one and
one-third as long as second (100 : 78), but shorter than first and with subparallel sides;
first and second recurrent veins both interstitial; cv-v straight, transverse, and coincident
with the point of origin of the basal vein. Hind wing with the transverse median vein
(cu-v) very oblique, only half as long as its distance from fork; jugal lobe short and
small, vannal lobe long, ending well beyond level of fork.

Integument throughout dull, except shining areas on summit of head, colour black
and dark brown, the posterior margins of abdominal tergites broadly pallid, and disk of
all tergites with distinct oily reflections. Punctation distinct, all punctures isolated,
moderately deep. Apex of tergite 7 of male with broadly rounded, distinctly undulated
margin (Fig. 94). Ventral surface of abdomen of male rather flat, both sexes with basal
part of first four sternites almost impunctate, postgradular areas closely punctate; all
posterior margins entire, but sternite 5 of male somewhat convex; sternite 6 of male
greatly modified, the apical angles with smooth, transverse tubercle ventrally (Fig. 95);
apodemes of sternite 7 of great length, directed laterad, lacking sclerotised transverse
ridges at the transition to disk, which is thin, membranous, exceptionally short and broad
(Fig. 96); sternite 8 of more normal shape, disk subrectangular (Fig. 97). Genital
capsule 1.5 mm long; intero-ventral process at apex of gonocoxite cylindrical, forming
a slender finger-shaped rod, subequal in length to gonostylus, which is much broader,
flat and spatulate, its apex squarely cut off (Fig. 100); penis valves of aberrant form,
the recurved subapical teeth unusually blunt, and distal portion tapering to a small,
bipartite knob (Fig. 98—99). Pygidial plate of female only partly visible, the end flat,
broadly triangular, with rounded apex.

Body vestiture much shorter and less dense than in most Habropoda, resembling
Elaphropoda much more closely. Occiput, genal area, thorax and basal segments of fore
legs posteriorly, clothed more densely with longish, raised, and heavily plumose hairs;
wide, triangular, median propodeal area with very short, decumbent radiating hairs.

Type species: Habropoda nubilipennis Cockerell.

Hab.: E China.

Habrophorula nubilipennis (Cockerell) comb. nov.

Habropoda nubilipennis Cockerell, 1930, Ann. Mag. Nat. Hist. (10) 6: 52 (9 Foochow, China).
— Lieftinck, 1966, Tijdschr. v. Ent. 109: 147 (notes).

Type material examined. — SE China, Fukien Prov.: 1 9 (holotype), with label
“Type, B.M. Type Hym. 17B.645” (printed on red-rimmed disk), “Habropoda nubili-
pennis Ckll. TYPE”, and “Foochow China (Kellogg)”, both in Cockerell’s handwriting
(BM).

Further material examined. — SE China, Fukien Prov.: 1 4 (diss, hitherto un-
described), Shaowu distr., Tachulan, 1000 m, 16—20.viii.1944, T. C. Maa (ML).

Original description:

“@. — Length about 13 mm.

Robust, black, with the aspect of Anthophora confusa, Sm.; eyes dark brown, inner
orbits parallel; clypeus coarsely punctured and obtusely carinate, the broad apical margin
and a narrower median stripe light yellow; a transverse but not triangular supraclypeal
mark, large triangular lateral face-marks (ending acutely on orbit at level of antennae),
large basal spot on the otherwise brown labrum, and basal half of mandibles, all light

M. A. LIEFTINCK: Review of Asiatic Habropoda 219

yellow; mandibles robust, curved, reddened beyond middle, and black and bidentate
at apex; hair of face very scanty, partly pale and partly fuscous, of front thin and pale
but some fuscous on upper part and vertex, but occiput with long pale fulvous hair;
lower part of cheeks with long white hair; cheeks above bounded by a keel, which is
continued across the top of head, behind the ocelli; antennae black, obscurely reddish
beneath; thorax densely covered with fulvous hair; tegulae light fulvous. Wings short,
dusky, shining, with dark nervures, the second cubital cell rather small, receiving first
recurrent nervure at end; the membrane is strongly glaucous. Legs dark reddish (not
ferruginous), with fulvous hair; hind spurs long and pale; abdomen finely punctured,
with a strong glaucous or bluish tint, tergites 2 to 4 with narrow cream-coloured bands;
sides of apex with long reddish-black hair; penultimate sternite fringed with red hair,
but the three before it with whitish hair.

Foochow district, China (Kellogg).

This cannot be the female of H. percarinata, the wings being quite different. It is
smaller than H. pekinensis, Ckll., with quite different abdominal bands.”

Female (holotype). — Apart from the sexual characters very nearly resembling the
male, described hereafter. Features worth mentioning in addition to those brought
forward in the original description and generic diagnosis, are the following:

Shape of head and face-marks, see Fig. 91; colours as in male. Labrum clear yellow,
a little longer than in male, gradually and slightly narrowed anteriorly. Punctation of
clypeus coarser, some punctures confluent and wider than interspaces. Length ratios of
antennal segments 3, 4 and 5 = 100:67:85 approx. Sculpture and pubescence of
head and thorax exactly as in male. Legs slender. Basitibial plate partly covered by a
hair tuft at apex of femur. Longitudinal ventral carina of hind tibia with thin hair-comb,
the surface smooth, shining and bare on distal half. Hind tibia and basitarsus as in
Fig. 93, the latter much shorter than tibia; surface of hind tibia well visible under the
long, strong brownish yellow scopal hairs, which are about as long as diameter of tibia.
Outer face of hind basitarsus with rather long, partly raised hairs shorter than on tibia
and nowhere hiding the surface; inner faces more densely pubescent, all hairs decumbent
and golden brown. Inner rami of all tarsal claws a little shorter than outer.

Abdomen as described for the male, the apical tergal fringes forming pale hair-lines,
the fringe at apex of tergite 5 not at all compact, and darker (golden-brown) only at
middle.

Male (hitherto undescribed; see also generic diagnosis). — Proboscis (fully extended)
7.2 mm, brownish-yellow; glossa densely hairy; galea smooth and hairless, 5.4 mm; first
segment of labial palpus almost as long as galea. Maxillary palpus slender, length ratios
of segments 1—6 as 18.2 : 100 : 54.5 : 82 : 59 : 22.7. Shape of head in frontal view and
face-marks as in Fig. 90, ground colour light chrome; teeth of mandibles obscured,
anterior border of labrum and labroclypeal suture ferruginous. Outer face of mandible
smooth, with few scattered punctures, labrum and clypeus finely chagreened, rather
shining, covered all over with large, shallow and partly flagging punctures, separated
by spaces equal to their own diameter; no distinct median ridge on clypeus; paraclypeal
area broad in proportion to clypeus, smooth and shining, impunctate, save for a few
scattered punctures at level of tentorial pits; frontoclypeal suture straight. Head other-
wise deep black; punctures circular and smaller than on face, those on summit of head
more widely spaced on smooth and shiny ground; surface of vertex, between ocelli and
upper orbits, distinctly impressed. Mouth-parts and face clothed sparsely with non-

220 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

LA pe

2S ek
COLA

=S
DIS:

Fig. 90—93. Habrophorula nubilipennis (Ckll.), structures, Fukien; Fig. 90—91, frontal view of
head of & (90) and the same, frontal and left lateral view of ® holotype (91); Fig. 92—93,
exterior view of left hind tibia-basitarsus of ¢ (92) and @ holotype (93)

plumose yellow hairs and erect bristles, the thin marginal fringe at labrum somewhat
longer; short, white, plumose hairs only around base of antennae, long, raised and
brown at occipital border, whitish and nowhere concealing surface on occiput and genal
area. Antenna brown; scape short and cylindrical, not fully twice as long as thick at apex
(10 : 6 approx.), not flattened anteriorly, but marked with an elongate yellow spot and
with sparse fringe of long pale hairs on either side; all remaining segments densely,
minutely tomentose; segment 3 less than half as long as 4 + 5, length ratios of first

M. A. LIEFTINCK: Review of Asiatic Habropoda 221

DAN)

yy,
Wy

Fig. 94—100. Habrophorula nubilipennis (Ckll.), & structures, Fukien; Fig. 94, ventral view of

seventh tergite; Fig. 95, exterior view of sixth sternite, showing apical tubercles; Fig. 96—97,

seventh (96) and eighth (97) sternites, exterior view; Fig. 98, genital capsule, ventral view; Fig. 99,

right gonoforceps and penis valves, slightly more detailed, dorsal view; Fig. 100, right gonocoxal
appendages, more enlarged, exterior view

six segments 100 : 23.7 : 37 : 50 : 58 : 60. Thorax black, dorsum and sides closely and
rather deeply punctate, punctures circular, separated by less than one puncture width;
propodeum almost vertical and flat, dark reddish-brown, surface densely and more finely
punctate, except a lozenge-shaped median area, which is smooth and impunctate.

222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974

Pubescence ochraceous, fading to greyish white laterally and underneath, all hairs con-
spicuously plumose but nowhere entirely concealing surface. Tegulae large, almost semi-
circular, brownish yellow, surface shining, very finely superficially punctate, partly
clothed with depressed, non-plumose hairs. Legs uniform yellowish brown; all segments
(except distitarsi) somewhat laterally compressed; femora slender, broadest at base,
tapering gradually, unarmed; tibiae slightly broadened apically; spurs yellowish, long
and almost straight, microscopically serrulate, those of hind legs of equal length; hind
tibia and basitarsus narrow (Fig. 92). Punctation of femora scanty and superficial, of
tibiae and tarsi somewhat more dense; pubescence short, hairs sparsely distributed,
except more crowded on the inside of basitarsi, though not quite concealing surface.
Abdomen elongate-oval, dullish, all pregradular areas of tergites and sternites dark
brown, the hind margins of 1—6 semitransparent, broadly yellowish and becoming
ferruginous at the transition; surface covered with small, evenly distributed, setiferous
punctures; all hairs brown, short and decumbent, except some longer raised ones dis-
persed at sides of tergites, which are paler in colour; also a single thin fringe of much
longer, golden yellow depressed hairs covering the pallid membranous apical borders
of the tergites. Postgradular areas of sternites smooth and shiny; pale anteapical fringes
not dense, increasingly longer posteriorly, longest at 5 and 6. Terminal segments and
genitalia, see generic diagnosis and Fig. 96—100.
Body length 12.0 mm approx., length of fore wing 8.0 mm.

The history of this highly remarkable bee is, briefly, as follows. The male and only
known specimen of that sex was discovered in a large collection of Anthophorini as-
sembled before and during the last world war by Dr. Tsing-chao Maa and his assistants
in the province of Fukien. This collection was entrusted to me for study and sent over
to Java as early as in 1953. It comprises a few thousand individuals belonging to four
closely interrelated genera, viz. Amegilla (sens. lat.), Anthophora, Elaphropoda and
Habropoda (sens. lat.). As the specimen in question was unique and on close inspection
proved to differ considerably in a number of characters from other constituents of the
tribe, it was set apart as something unfamiliar and left unattended for more than twenty
years. As pointed out already in my preliminary survey of Old World anthophorines
(Lieftinck, 1966), it would become necessary sooner or later to investigate more fully
the Asiatic members of Habropoda and its immediate allies, several species requiring
further study. The outcome of this can be found in the present paper. One of the critical
taxa to be dealt with was Habropoda nubilipennis Ckll., a species already briefly com-
mented upon in my former account (loc. cit.: 147). It was described from a single female
collected in Fukien, but no suitable partner for it had ever turned up. The male being
still unknown at the time of characterizing the female, Cockerell compared nubilipennis
with another member of the genus which he described in the same paper as Habropoda
percarinata. This was a male collected in Fukien at the same locality with the former.
Though Cockerell observed that it resembled nubilipennis somewhat in general appear-
ance, he nevertheless recognized it as a distinct species. Later investigations confirmed
this view but at the same time the insect soon proved to be a member of the new genus
Elaphropoda, both sexes of E. percarinata (Ckll.) being well represented in Dr. Maa’s
collections (Lieftinck, loc. cit.: 157—160). In the article cited above I left the female of
Habropoda nubilipennis as it stood, and it was not until last year that I could re-examine
the type of Cockerell’s bee in the collection of the British Museum (Nat. Hist.). This
was directly confronted with the solitary male from Fukien, a proceeding which enabled

M. A. LIEFTINCK: Review of Asiatic Habropoda 223

me at last to establish the true identity of the latter. It means that a careful study of all
salient characters revealed beyond doubt that the two are not only conspecific but should
be transferred to a new genus, here defined under the name of Habrophorula, the type
species consequently being Habropoda nubilipennis Cockerell.

The male of H. nubilipennis is immediately distinguished from all other EDER
of the tribe occurring in the same region by the unique form of the terminal abdominal
segments and genital organs, especially the shape of the hidden sternal plates and separate
parts of the genital capsule being totally different from anything seen in allied genera.

On comparing Habrophorula with other genera in which the first recurrent nervure
and distal side of the second submarginal cell in the fore wing are interstitial (or nearly
so), the genus can be recognized by the following combination of characters: (1) body
form markedly elongate; (2) labrum small with simply convex, projecting anterior
border; (3) proboscis long and galea attenuated; (4) mandibles unidentate; (5) clypeus
short and very little protruding; (6) ocelli approximated, forming together an isosceles
triangle; (7) antennal segment 3 abbreviated and much shorter than succeeding segments;
(8) no long hair tufts on vertex in front of and just behind anterior ocellus or between
the latter and inner orbits; (9) legs extremely slender and unmodified (4 9) and
scopal hairs thin (£); (10) marginal cell on fore wing of great proportional length,
proximal and distal sides of 3rd s.m. weakly convex and almost parallel; transverse
median nervure in hind wing very oblique, much shorter than its distance from fork;
(11) dense silky hair covering thoracic segments conspicuously plumose; and (12) fine
decumbent hairs covering disks of gastral tergites rather sparse and notably short, coupled
with pale translucent posterior tergal margins and absence of conspicuous posterior
sternal hair fringes.

The extraordinary array of characters here described and illustrated suffices to
distinguish the new genus from Habropoda. In stature and some peculiarities of the
venation and vestiture Habrophorula is rather strongly suggestive of Elaphropoda,
sharing with this the characters mentioned above sub 1—3, 6, 8, 9 (female only), 10
(partly), 11 and 12. There are important differences, however, particularly the non-
protuberant face, different proportional lengths of the proximal antennal segments, and
the simple slender legs, are features which will serve to the easy recognition of Habro-
phorula, to say nothing of the very unusual structure of the male terminalia. Additional
distinctives of Elaphropoda precluding the possibility of any close relationship between
the two, are: (1) antennal segment 4 (instead of 3!) annular and much shorter than
either 3 and 5; (2) mandibles bidentate interiorly; (3) proximal and distal sides of 3rd
s.m. not at all parallel, the cell itself being much shorter than in Habrophorula.

In connection with the above it is of interest to mention the recently proposed Central
American genus Deltoptila LaBerge & Michener, 1963 (type species Habropoda monte-
zumia Smith), also of the Anthophorini. This was established some years in advance
of Elaphropoda Lieft., from the eastern hemisphere, sharing with this the enormously
protuberant clypeus, great length of both the proboscis and marginal cell of the fore
wing, and the somewhat similarly formed male terminalia. For important differences
between the two, see the existing descriptions.

Note. — In 1966, at the Paris Museum, I have cursorily examined a series of both
sexes of the Mexican Anthophora badia Dours, 1869, from Oaxaca in Sichel’s col-
lection. This bee was not included in the specific list given by LaBerge & Michener,
but undoubtedly also belongs to Deltoptila.

224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 5, 1974
REFERENCES

Cockerell, T. D. A., 1905. — The bees of Southern California. — Bull. South. Calif. Acad. Sci. 4:
99—106.

Dours, J. A., 1869. — Monographie iconographique du genre Anthophora Lat. — Mém. Soc.

linnéenne Nord France (Amiens), 2: 1—211, 2 pls.

LaBerge, W. E. & C. D. Michener, 1963. — Deltoptila, a Middle American genus of anthophorine

bees (Hym., Apoidea). — Bull. Univ. Nebraska State Mus. 4 (9): 209—225, 26 fig.
Lieftinck, M. A., 1966. — Notes on Anthophorine bees, mainly from the Old World (Apoidea). —
Tijdschr. v. Ent. 109: 125—161, fig. 1—92.
, 1972. — Further studies on Old World melectine bees, with stray notes on their distribu-
tion and host relationships (Hymenoptera, Anthophoridae). — Tijdschr. v. Ent. 115: 253—
324, tig. 1-55, pl 1-2:
Michener, C. D., 1944. — Comparative external morphology, phylogeny, and a classification of the
bees (Hymenoptera). — Bull. Amer. Mus. Nat. Hist. 82: 151—326, tfig. 1—246, diagr.
1-13.

Mitchell, T. B., 1962. — Bees of the Eastern United States, II. — Tech. Bull. 152 North Carolina
Agric. Exper. Station, 557 pp., 134 fig., 18 tab.

Popov, V. B., 1948. — Geographical distribution of Apidae of the genus Habropoda F. Smith. —
Dokl. Akad. Nauk USSR, new ser. 59: 1673—1676, 1 map (in Russian).

Timberlake, P. H., 1962. — An interesting new species of Emphoropsis from California. — Ent.
News, 73: 38—39.

INDEX

Unless indicated otherwise specific names are of Habropoda. Synonyms are in italics.

alashanica 179 magrettii, Elaphropoda 186
maiella 193

mendax, Bombus 183
mimetica 164, 174 (keys), 183
moelleri, Elaphropoda 186
montana 186, 205

badia, Deltoptila 223 montezumia, Deltoptila 223

bizonatus, Bombus 183 nubilipennis, Habrophorula 218
bucconis 167, 178 (keys), 201

Amegilla 180, 222
Anthophora 212, 222
apatelia 171, 178 (keys), 215
apostasia 168 (key), 196

orbifrons 173 (key), 182

pekinensis 163, 172 (keys), 179
pelmata 169 (key), 210
percarinata, Elaphropoda 219
plantifera 170 (key), 208

defector (ssp.), Bombus 183
deiopea 171, 178 (keys), 212
Deltoptila 223

disconota 175 (key), 185

Elaphropoda 186, 222 pseudobomboides, Anthophora 183
Emphoropsis 159, 180 radoszkowskii 169, 177 (keys), 205
excellens, Emphoropsis 160 rowlandi 166, 175 (keys), 188
fulvipes, Elaphropoda 186 sinensis 164, 173 (keys), 180

sutepensis 168, 175 (keys), 198
Habrophorula 158, 217

habropodae, Protomelissa 199 tainanicola 166, 176 (keys), 190
hookeri 177 (key), 199 tarsata 186

MUTA tricolor, Protomelissa 188
imitatrix 170, 178 (keys), 203 tumidifrons 173 (key), 182
krishna 216 turneri 165, 174 (keys), 185

laboriosa, Emphoropsis 160 zonatula 179

" ar a

- No. 4. F. Willemse, 1968. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov
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B No. 5° €. A. W. Jeekel, 1970. — Nomenclator generum et familiarum. Diplopodorum, 412 pp.
| D.FI. 100.—.

No. 6. J. G. Betrem, 1971. — The African Campsomerinae (Hymenoptera, Scoliidae), 326 PP..
| 47 figs., 6 pls. D.Fl. 120.—.

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Ta RON

SYSTEMATICS AND EVOLUTION OF THE
PALAEARCTIC SPIALIA SPECIES (LEPIDOPTERA,
HESPERIIDAE)

by
R. DE JONG

Rijksmuseum van Natuurlijke Historie, Leiden

ABSTRACT

The Palaearctic members of Spialia have been revised taxonomically. Two species groups have been
distinguished, viz. the phlomidis group and the sertorius group. The former comprises the species
phlomidis, osthelderi, geron and doris, and is related to the African species spo. The sertorius group
has two Palaearctic members, viz. sertorius and orbifer, united in a superspecies, and two members
outside the Palaearctic, viz. the African mafa and the Indian galba.

By analyzing some characters as to their plesiomorphous or apomorphous conditions, a reconstruc-
tion has been made of the evolutionary and geographic history of the two species groups. This
reconstruction forms the theoretical basis for the arrangement of the species and subspecies adopted.

CONTENTS
OL GO nei di i. re ZO
2, Geelen LEE EL ENE de à ee LC
è Vigilante ee Oe NT,
BNelkhowledsements MEERE REED AI) Le NER DD
dh ic:hodsyandmeasurements aan nme. en oder le ed 22
Zee DElimitattonvand classification’ of the genus. Spialiai . | . an NN P28
i. Ilie Pie group: ME a ea
eI CE LO SPECIESEPIOUPL IT ne © sl e DA
5. Evolutionary and geographic history of the genus Spialia in the Palaearctic region. . 256
Le CERA IRR EE AMAT
EREN VOUS SOUP RARE OO MO oe ZG
MIEVOlutionary history: MINI ER BRAMA i Z.ZT
PAMGcopraphicchistoryi de: loan DS a Le dina re Pe eg
TEE 20271, CLOUD l'al AR e ale ra 1260
iaBEyolubiovary, history a EN neer Zee 2. ZOO
PiGcosraphico history rage e. n PE 263
di STE EE SME ee Geo 267
U AT vot Lomé 4 APM: 25 nea Sia Kee | 268

225

226 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

1. INTRODUCTION
1.a. General

The aim of this study is to examine the phylogenetic relationships of the Palaearctic
members of the genus Spialia by means of a reconstruction of the evolutionary and
geographic history. For this purpose the species concerned have been revised taxon-
omically and several characters have been analyzed as to their conditions being derived
or primitive. This leads to an arrangement of the species supposedly reflecting the phylo-
genetic relationships. This arrangement is not very different from former arrangements,
but especially in sertorius and its allies the solution of the perpetual problem “species
or subspecies?’ is somewhat different from all former solutions. This is mainly due to
the subspecies concept.

Under the term “subspecies” usually a number of various types of geographic variation
is described. This fact appears to be little recognized and virtually is a source of much
confusion. Some authors apply the term “subspecies” to any recognizable population,
others use the term to denote rather arbitrary parts of the geographic variation. Recently,
Dobzhansky (1970) defined a subspecies as “a race that a taxonomist regards as suf-
ficiently different from other races to bestow upon it a Latin name”. To my opinion
such a concept must lead to confusion. This is not the place for an exhaustive discussion
on the subspecies concept, therefore I will only stress that for a student wishing to trace
the geographic history of a species, only that part of the geographic variation is useful
for him which arose by geographic isolation. Consequently, in my paper on the genus
Pyrgus (De Jong, 1972) I presented the following definition:

A subspecies is an, at present or formerly, geographically isolated group of populations
of a species, which can be distinguished by one or more characters from other such groups
of populations of the species.

The fact that the existence of a former geographic isolation must remain hypothetical
cannot be a serious objection, so long we do not use the subspecies thus defined for
proving the former existence of such an isolation. Virtually the distinction of subspecies
is a hypothesis about the geographic history of the species. The hypothesis may be wrong,
but the meaning is clear. I cannot see the use of a subspecies that is an aggregate of
various kinds of variation. Some people may object that we better stick to the facts, but
what is the use of facts we do not use?

The remainder of the geographic variation lays in the field of the ecologist and
geneticist, rather than in that of the zoogeographer. However, the last-named is in the
best position to notice this variation. Without experiments it is difficult to ascertain
whether a variation is hereditary or not. Nevertheless, I have attempted to distinguish
between non-genetic and genetic variation, which I have termed ecophenotypic and clinal,
respectively. I consider it senseless to use a trinomen in the case of clinal variation. It not
only would cause confusion with subspecific distinctions, but it also would suggest dis-
continuities where there are only continuities. It seems better to describe clinal variation
than to denominate it. In many cases, however, parts of the clinal variation have already
been described as subspecies. In such cases I am using Latin names, but separately and
not as a trinomen.

The same applies to ecophenotypic variation. A large part of the nomenclatural and
taxonomic confusion, at least in butterflies, is due to the practice of describing ecopheno-

R. DE JONG: Systematics and evolution of Spialia 227

typic varieties as subspecies (e.g. the form Azbzscae of sertorius, see chapter 4).

b. Wing markings and genitalia

For the wing markings and genitalia I may refer to my paper on Pyrgus (De Jong,
1972). However, some parts of the male genitalia of Spialia are not so highly dif-
ferentiated as in Palaearctic Pyrgus species. Therefore, the names of the parts can more
easily be brought into accordance with the more currently used names as expressed in
Tuxen (1970). So, with regard to the names used in Pyrgus the following alterations
are made in this paper:

(1) lateral apophyses become gnathos. In Spialia this structure is much simpler than
in Pyrgus.

(2) harpe becomes costa. In Pyrgus the distal end of the dorsal part of the valve is
highly differentiated. Also in Spialia this part is differentiated, but in a different way
and presumably by an independent evolution. It seems advisable to term this structure
in Spialia by a trivial name and I use the name costal process. It would be senseless
to replace the terms style, stylifer and antistyle in Pyrgus also by costal process.

(3) cuiller becomes cucullus. Also in Pyrgus it may be advisable to introduce this
alteration.

c. Acknowledgements

It is a pleasure for me to express my gratitude to the following persons and institutions
for offering me the opportunity of studying the material in their collections or for
presenting specimens (abbreviations used in the text are placed in brackets):

W. L. Blom (Groningen) (WLB), H. Coene (Amsterdam) (HC), Dr. W. Forster
(Zoologische Sammlung des Bayerischen Staates, Munich) (ZSM), Dr. L. Gozmany
(Hungarian Natural History Museum, Budapest) (MB), T. G. Howarth (British
Museum, Natural History, London) (BM), Instituut voor Taxonomische Zodlogie
(Zoölogisch Museum, Amsterdam) (ITZ), Mrs. A. Vartian and Dr. F. Kasy (Natur-
historisches Museum, Vienna) (MW), J. H. Lourens (Amsterdam), H. van Oorschot
(Amsterdam) (HO), J. Wolschrijn (Heerde, Netherlands) and C. W. Wyatt (Farnham,
England) (CW). The Rijksmuseum van Natuurlijke Historie (Leiden) is abbreviated:
(ML).

d. Methods and measurements

All names given to species, subspecies and forms of Spialia in the Palaearctic have
been included in this study, with the exception of the few names relating to individual
varieties. A full account of these names can be found in Evans (1949).

To avoid confusion between species and subspecies in the text, a species is indicated
by the combinations Spzalia a-us, S. a-us, or simply 4-5, a subspecies by the combination
a-us b-us ot by ssp. b-us. Names referring to the infrasubspecific variation can be rec-
ognized by the word “forma” preceding the name of the variety.

Measurements refer to the length of the fore wing from base to apex.

228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

2. Delimitation and classification of the genus Spzalia

The genus Spzalia was erected by Swinhoe (1913 : 99) to include the species galba
(type-species), zebra, geron and sao which were previously placed in the collective genus
of “black and white Skippers” known under the names Hesperia and Syrichtus. Thus,
Swinhoe only included species occurring within the boundaries of the former British
India. Evans (1937) placed also the Ethiopian black and white Skippers in Spialia. This
extension of the genus is generally accepted, also in this study.

As a revision of the generic divisions of the Pyrgus group (Evans, 1949) is intended
in the near future, it may suffice now to mention only the most conspicuous characters of
Spialia, as against the genera Pyrgus and Muschampia (sensu Evans; recte Syrichtus).
The fact that the last named genus is heterogeneous does not need to bother us here.

From Pyrgus species those of Spialia can easily be distinguished by the presence of a
more or less well-marked and complete series of submarginal spots on the fore wing
upperside and by the place of the median spots in spaces 4 and 5 (if present) that are
in line with the median spots in spaces 6 to 8. In Pyrgzs the submarginal spots are
incomplete and faint or absent and the median spots in 4 and 5 (usually present and
distinct), are not contiguous to the spots in 6 to 8, but much nearer the termen.

Other distinguishing characters can be found in the males, viz., a costal fold in the
fore wing and a hair tuft on the hind tibiae, fitting into a thoracic pouch in Pyrgus.
Both these secondary sexual characters are absent in Spialia, only the males of some
Ethiopian species have an indistinct costal fold.

From Syrichtus species Spialia species can be distinguished by the median spot in space
2 of the fore wing upperside which is central between the cell spot and the median spot
in space 3 or nearer the latter. In Syrichtzs species the median spot in 2 is nearer to the
cell spot than to the median spot in 3. Moreover, the termen of the hind wing is more
or less crenulate in various Syrichtus species, but never so in those of Spialia. The males
of Syrichtus species usually have a costal fold.

Few authors have tried to subdivide the genus Spzalia. Warren (1926), while dealing
with Palaearctic species (under the name Powellia Tutt, a junior homonym of Powellia
Maskell, 1879; see Verity, 1940), distinguished two species groups, viz. the geron group
and the serforius group. I disagree with Warren about the assignment of species to these
groups.

Picard (1947) proposed a more radical subdivision by erecting a new genus, viz.
Platygnathia for phlomidis, geron and doris. However, he did not study the Ethiopian
species and therefore, his classification is somewhat premature. For a revisional study of
the taxonomy of the genus which I hope to complete in the near future, I have studied
the male genitalia of all Ethiopian Spialia species. As a result of this study I consider
the species spo a link between the relatives of phlomidis and the Ethiopian species. The
genus may be subdivided into species groups, but there is no apparent use in making
genera of such groups. I am an opponent of the use of subgenera: their application does
not solve problems, but only adds to nomenclatural difficulties. Therefore I propose to
subdivide the genus Spialia only into species groups, while placing Platygnathia as a
junior subjective synonym of Spialia.

In the Palaearctic two species groups can be distinguished:

(1) the phlomidis species group, including the species phlomidis, geron, osthelderi
and doris.

R. DE JONG : Systematics and evolution of Spialia 229

Fig. 1—8. Right valva and aedeagus of Spialia. 1—2, S. phlomidis phlomidis (Amasia); 3, S.

geron struvei (Ferghana); 4—5, S. geron geron (Baluchistan, Kahan); 6, S. osthelderi gecko

(Hyrcania); 7—8, S. doris daphne (S. Morocco, Ksar-es-Souk). Valva seen from the inside, aedeagus
seen dorso-laterally from the left side

Male genitalia (Fig. 1—8). — Aedeagus with a strongly sclerotized and indented
crest, latero-ventral in the middle, or a long, unindented branch in the same place, and
with an extended apex. Cucullus spined at apex. Costa spined on dorsal surface and at
the ventro-distal end, or only at the ventro-distal end.

Female genitalia (Fig. 9—13). — Two heavily sclerotized and indented wing-like
structures at the base of the eighth sternite. No distinct genital plate.
External characters. — On the upperside of the fore wing a conspicuous spot or bai

at the end of the cell.

230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

Fig. 9— 11. Female genitalia of Spialia. 9, S. phlomidis phlomidis (Greece), ventral side; 10,
idem, seen from the left; 11, S. geron geron (Baluchistan, Kahan), ventral side

(2) the sertorius species group, including the species sertorius and orbifer (united
into a superspecies). Two non-Palaearctic Spialia species can also be referred to this
group, viz. the Oriental galba and the Ethiopian mafa.

Male genitalia (Fig. 24—35). — Aedeagus without crest or branch and apex not
extended. Cucullus little differentiated, without spines. Costa at ventro-distal end with
a more or less ellipsoid extension bearing long upcurved spines. Between costa and
cucullus a densely haired fold at the inside of the valva.

Female genitalia (Fig. 21—23). — No wing-like structures. In Palaearctic species
genital plate narrow in proximal half, well developed.
External characters. — On the upperside of the fore wing, spot at end of cell faint

or absent in Palaearctic species; galba can be distinguished from the phlomidis group by
the inner spot in space 7 of the hind wing underside being placed directly over the basal
spot in the cell; mafa has a pale basal spot in space Ic of the hind wing underside,
which is absent in the phlomidis group.

R. DE JONG: Systematics and evolution of Spialia 231

Fig. 12—13. Female genitalia of Spialia, ventral side. 12, S. osthelderi osthelderi (Akbès); 13,
S. doris doris (Aden)

3. THE phlomidis SPECIES GROUP

The species of the phlomidis group can be regarded as closely related and Evans (1956:
750) is correct in stating that they “looked at from a very broad point of view, might be
treated as conspecific’. As a great overlap exists in the distributional areas of the species,
it is undesirable to unite them even into a superspecies, but by bringing them together
into a species group we can indicate their supposedly close relationship.

The species can be distinguished as follows.

Key to the species of the phlomidis group

1. Central band of hind wing underside directed and more or less conjoined to inner
spot in space 7 (Fig. 18). Costa of valva without spines dorsally (Fig. 1). Aedeagus
with a sclerotized crest medially (Fig. 2). Papillae anales (Fig. 14) rather pointed,

232

Fig.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

basal lobe large, covering about three of base; length ratio papilla : apophysis
posterior we I SR NES fas AI
Central band of hind wing Hide not toen to inner spot in space 7, but
directed more or less to apex. Costa of valva with spines dorsally. Aedeagus with a
sclerotized crest or branch medially. Papillae anales bluntly rounded, basal lobe
covering half of base or less, apophyses posteriores relatively shorter. . . . 2
Central band of hind wing underside more or less directed to outer spot in space 7,
without touching it (Fig. 20). Discal spot in space Ib of fore wing upperside not
touching vein 1. Small, & 8.6—12.4 mm. Costa of valva distally strongly concave
and reaching beyond cucullus (Fig. 7). Aedeagus with a sclerotized crest medially
(Fig. 8). Basal lobe of papillae anales (Fig. 17), a small flap, nearly circular; length
ratio papilla : apophysis posterior, 5:75. . . a RO TES
Central band of hind wing underside directed to a aati en the inner and outer
spots in space 7 (Fig. 19). Discal spot in space Ib of fore wing upperside usually
touching vein 1. Costa of valva not extending beyond cucullus. Basal lobe of papillae
andles elongate‘-TLärser, MOMENT

14 17

14—17. Papillae anales of Spialia. 14, S. phlomidis phlomidis (Greece); 15, S. geron geron
(Baluchistan, Kahan); 16, S. osthelderi gecko (Kerman); 17, S. doris doris (Ras Farták)

Colour of hind wing underside greenish. Spot at base of cell of fore wing upperside
and cell spot of hind wing upperside present, or if absent, then colour of hind wing
underside whitish with faint markings. g' 11—13.6 mm. Costa of valva strongly
curved (Fig. 6). Aedeagus with a crest medially. Papillae anales with short apophyses
posteriores; length ratio papilla : apophysis posterior, 5 : 5—6; basal lobe broad

R. DE JONG: Systematics and evolution of Spialia 233

IO a 4 dt) Morhelder
— Colour of hind wing unddersicie dir El zi, ee ci fore and hind wing
uppersides usually absent. $ 11.7—14.2 mm. Costa of valva more gently curved
(Fig. 4) *). Aedeagus with a branch medially (Fig. 5). Papillae anales (Fig. 15)
with slightly longer apophyses ee a ratio a : apophysis posterior,
5 : 7, basal lobe narrower . $ : «Mal verte et achog

18 19 20

Fig. 18—20. Underside of left hind wing of Spialia. 18, S. phlomidis phlomidis (Greece); 19
S. osthelderi gecko (Semnan); 20, S. doris evanida (Karachi)

Note. The differences in the female genitalia are much less conspicuous than in those
of the male. It is recommendable to examine the papillae anales freely floating in water
or alcohol, as their shape is easily influenced by the pressure of a cover glass.

There are also differences in the sclerotization of the postvaginal region. This is evenly
sclerotized in phlomidis, with a central narrow membranous gap in osthelderi and doris,
and with a wide central membranous region in geron.

The strongest sclerotization of the wing-like structures was found in osthelderi and the
weakest in doris. However, this may be partly due to differences in mounting (e.g. length
of maceration by KOH).

Spialia phlomidis (Herrich-Schäffer)

Distribution (Map 1). — From Macedonia and Albania through Greece and Turkey
to NE Iran. A local species, known only from the following localities:
Albania : Kula e Lumes, Maliqsee, Korce (Alberti, 1965; material examined);
Yugoslavian Macedonia : Orasje, Drenovo, Ohrid, Petrina planina (Thurner, 1964;
material examined) ;
Greece : Morea (Rebel, 1902); further only “Greece” (Oberthür, 1912; material
examined) ;
Turkey : Brussa, Amasia, Tokat, Goynucek, Ayac Dagi, Berud Dagh (Taurus),
“Armenia” (Staudinger, 1881; Oberthür, 1912; material examined);
South Russia: “S. Russia”, “Caucasus”, Kasikaporan, Ordubad (material examined) ;
Iran : Shahrud, “Hyrcania” (material Er mined):
Syria : Akbès (Oberthür, 1912);
Lebanon : Bludan (Ellison & Wiltshire, 1939), Mt. Hermon (material examined).

*) The figure of the valva of geron by Reverdin (1914) represents that of osthelderi!

234 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

Groum-Grshimailo (1890) recorded this species from Pamir, but this observation
probably concerns osthelderi gecko or geron struvei.

Habitat. — According to Thurner (1964) in Macedonia mainly at low altitudes,
exceptionally up to 1600 m; in Turkey to 1300 m (Ayac Dagi, material examined), at
Bludan between 1700 and 2000 m (Ellison & Wiltshire, 1939). Always in dry and hot
localities.

Biology. — All the known specimens are from June and July. No further records
about the biology.
Geographic variation. — Two subspecies can be distinguished. The material, however,

is very scanty, especially from the eastern part of the range.
Spialia phlomidis phlomidis (Herrich-Schäffer)

Herrich-Schäffer, 1845. — Syst. Schmett. Eur. 1 : 153; Hesp. pl. 2 fig. 8, 9. (For date of publica-
tion, see Hemming, 1937).
Type-locality: Turkey (shores of the Sea of Marmora).

= = T T Wa T DI

CR IS) È
>
I NW N\
CxS dd da
Temi Se Tare
® Ta
u
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) à Map 1 - Spialia phlomidis

ei = À 1 RI =

Material examined. — 105 g' 33 9 : 28 S 10 9 Macedonia (Ohrid) (HC, ML,
ITZ), 21 & 10 9 Albania (Korce) (BM), 12 & 2 Q Greece (BM, ML), 37
d 9 9 Turkey (Brussa, Tokat, Amasia, Goynucek, Ayac Dagi, Berud Dagh, “Pontus”,
Turkey’) (BM, ML), 1 g 1 2 Kasikaporan, 2 g' Caucasus, 2 9 S. Russia, 1
d' Ordubad, 1 & 1 9 Iran (Shahrud, “Hyrcania”) (all BM).

Distribution. — As that of the species, except Anti-Lebanon and Mt. Hermon.

Subspecific characters. — g' 13.2—15.7 mm. A large and strongly marked form, if
compared with ssp. hermona.

Variation. — Hemming (1932a) compared a long series of freshly emerged specimens
(mainly males) from Mt. Chelmos and two specimens from Parnassos with specimens
from E. Turkey, Armenia and Iran. He found differences, the specimens from the eastern
part of the range being slightly larger, greyer and with larger spots. On account of this
difference he described a new subspecies, viz. phlomidis eupator (type-locality: Amasia),

VT SE

O oe Die

R. DE JONG: Systematics and evolution of Spialia 235

occurring in the eastern part of the range of phlomidis, while phlomidis phlomidis flies
in the Balkans and in the western part of Asiatic Turkey (e.g. at Brussa).

I could examine much more material from the Balkans than Hemming. This material
does not support the separation made by Hemming. Although the holotype and allotype
of phlomidis eupator (in the BM) agree very well with the description, they are not
representative for the eastern populations, as far as can be judged from the limited
number of specimens available. We have to wait for further material from the eastern
part of the range of phlomidis before we can hope to make a subspecific separation.

Spialia phlomidis hermona Evans

Evans, 1956. — Ann. Mag. Nat. Hist. 9 (12) : 750.
Type-locality: Mt. Hermon.

Material examined. — 1 g° Mt. Hermon, 30.VII.1945 (holotype) (BM).

Distribution. — Only the type specimen is known, but Ellison & Wiltshire (1939)
recorded the occurrence of a small phlomidis near Bludan in the Anti-Lebanon, that
undoubtedly concerns this subspecies.

Subspecific characters. — According to Evans (1956) this subspecies is smaller than
phlomidis phlomidis, 3° 12 mm, and on the upperside exactly like osthelderi gecko, but
the valva resembles phlomidis phlomidis. To this I can only add that the fringes seem
to be somewhat narrower relatively than in phlomidis phlomidis and that I was not able
to find an essential difference in the male genitalia.

Spialia osthelderi (Pfeiffer)

Distribution (Map 2). — From SE. Turkey to Central Asia, but only few localities
are known:

Turkey: Marash, Diarbekr (Pfeiffer, 1932; material examined); Sivrice (De Lattin,
1950); Yüksek Dagh (Amanus Mts) (material examined).

Syria: Akbès (material examined).

Lebanon: Bscherre (material examined).

Iraq: Kizil-Robat, Mirjana, Suleimanyeh (material examined).

Iran: “Hyrcania”, Shahrud, Keredj, Semnan, Mashad, Seguck near Kerman (material
examined); Shiraz, Sine-Sefid, Tchouroum (Brandt, 1939; material examined).

Afghanistan: Reschke (material examined; I was not able to trace this locality,
possibly this is not the name of a locality but of a collector). Evans (1949) recorded
2 d 1 9 from Kabul; these specimens, however, proved to belong to geron.

C. Asia: Ketmen Tjube (Sussamyr Mts) (material examined).

Habitat. — According to Pfeiffer (1932) flying on slopes up to 1000 m; in SW Iran
up to 2600 m (material examined); in Iraq desert foothills and lower middle heights
of the mountains (Wiltshire, 1957).

Biology. — According to Pfeiffer (1939) two generations per year. The specimens
examined have been collected in all months from April till July and in September.

Geographic variation. — Evans (1949) distinguished two subspecies as follows:

(1) ssp. ostheldert — Above markings reduced, particularly on the hind wing upper-

236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

side, where the discal spot in space lc and the basal cell spot are absent usually. Hind
wing underside almost all white, with all markings faint.

(2) ssp. gecko — Above and below with well-marked spots. On the upperside of the
fore wing basal cell spot present, as also the cell spot on the hind wing upperside.

This suggests a clear-cut difference between the two subspecies, but the impression
is wrong. The only difference upon the upperside I could establish concerns the discal
spot in space Ib of the hind wing which is always visible and sometimes large in ssp.
gecko, mostly inconspicuous or absent, but sometimes also large (e.g. G' paratype),
in ssp. ostheldert.

Also on the underside of the hind wing the difference is not clear-cut. In spp. osthelderi
the ground colour is always whitish, so that the white markings are very faint. In ssp.
gecko the ground colour is olive greenish or brownish with well-marked spots, but in
two males from the vicinity of Shiraz (Comée, 2600 m; Sine-Sefid, 2200 m) the ground
colour is whitish and the markings faint, so that these specimens are indistinguishable
from ssp. osthelderi.

As the material is so scanty, the Erlen of differentiating characters needs not be
significant, but it can also mean that there is a large transitional zone or a clinal variation
in the character mentioned. For the time being it seems advisable to consider both forms
separate subspecies. The white underside of the hind wing does not appear to be a
character of a purely ecological nature, forming a part of a cline. It is interesting to note
that this character also occurs in the two Hesperid species Pyrgus melotis melotis
Duponchel (see De Jong, 1972) and Syrichtus tessellum nomas Lederer (see Warren,
1926), both occurring in Turkey and Syria, much like osthelderi osthelder:.

Spialia osthelderi osthelderi (Pfeiffer)

Pfeiffer, 1932. — Mitt. Münch. Entom. Ges. 22:50, pl. 4 fig. 14, 15, 18.
Type-locality: Marash (Turkey).

Material examined. — 9 8 6 9 : 4 gd 2 9 Marash (3 G 2 Q paratypes) (ZSM,
BM), 1 8 Diarbekr, 1 & Amanus (Yüksek Dagh), 1 8 Bscherre (Lebanon) (all
ZSM), 1 9 Akbès, 1 g 3 $ River Dyala (Kizil-Robat, Mirjana), 1 G' Suleimanyeh
(Kurdistan) (all BM).

Subspecific characters. — See above. g' 11.5—13.6 mm. Small specimens possibly
belong to a second generation.

Distribution. — The western part of the range, eastward probably to W. Iran.

Spialia osthelderi gecko Evans

Evans, 1949. — Cat. Hesp. Eur. Asia & Austr. : 176.
Type-locality: Hyrcania.

Nomenclature. — Pfeiffer (1939) mentioned a new form of osthelderi that he would
name and describe before long. Shortly thereafter, in a list of butterflies from S. Iran,
Brandt (1939) mentioned “Spialia osthelderi struveoides Pfeiffer”. This is undoubtedly
the same form as gecko, since gecko looks somewhat like Spialia geron struvei (at least,
if compared with osthelderi). As the name siruveoides was published in 1939, gecko
would fall as a junior subjective synonym. However, Dr. Forster kindly informed me

R. DE JONG: Systematics and evolution of Spialia 237

that Pfeiffer never published the name struveoides. As the name is mentioned by Brandt
without a description, it is not available in nomenclatural sense.

core T =a AT

CR =
EIS
da Ei

5 è N deo @ \ 5

ID 0
oO
Y !
N 3; Map 2 - Spialia osthelderi

Ô 8; Map 3 - Spialia geron

Material examined. — 15 g 5 9 :4 g 1 9 Hyrcania (1 G' holotype) (BM), 1 ¢
Semnan (MW), 1 & Mashad (WLB), 3 g' Keredj, 5 gd 1 9 Fars (Shiraz, Sine
Sefid, Tchouroum, Comèe) (all ZSM), 1 g' Shahrud, 1 9 Seguck near Kerman, 1 9
Afghanistan (Reschke), 1 9 Ketmen Tjube (Sussamyr Mts) (all BM).

Distribution. — The eastern part of the range, from Iran to C. Asia.

Subspecific characters. — See above. gi 11—13.6 mm. Small specimens may belong
to a second generation.

238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

Spialia geron (Watson)

Distribution (Map 3). — From Ordubad (Russian-Iranian frontier, west of the
Caspian Sea) to Barkul and Hami (eastern end of Tian Shan), but extremely local and
only known from the following localities:

Iran: Ordubad (Evans, 1949: “Ordub’’) (material examined).

Afghanistan: Arbarp (10 mls W. of Kabul) (material examined; wrongly identified
as osthelderi by Evans, 1949).

W. Pakistan: Baluchistan (Quetta and surroundings) (Watson, 1893; material
examined).

C. Asia: ‘“Ferghana” (material examined); Issyk Kul, Barkul (Püngeler, 1914);
Dsharkent (= Panfilov) (Pfeiffer, 1932; material examined); Hami (material exam-
ined).

The Pyrgus phlomidis recorded by Groum Grshimailo (1890) from Pamir, probably
is the present species, or osthelderi gecko. Clench & Shoumatoff (1956) listed a badly
worn female from Panjao, southwest of Koh-i-Baba (Afghanistan) as a possible geron.
This identification may be correct, but the specimen may also belong to osthelderi gecko.

Habitat. — Not recorded.

Biology. — Presumably two generations per year. The specimens from Baluchistan in
the BM date from the periods February to March, and May to September, respectively.

Geographic variation. — The specimens from Afghanistan and Baluchistan differ
constantly in size and in the male genitalia dissected: ten males from Afghanistan and
Baluchistan, four males from Ferghana and one male from Ordubad.

Spialia geron geron (Watson)

Watson, 1893. — Proc. Zool. Soc. London 1893: 66.
Type-locality: Quetta (Baluchistan).

Material examined. — 42 g' 20 9 :2 & 3 Q Arbarp (Afghanistan, 10 miles W. of
Kabul), 40 & 17 9 Baluchistan (Quetta, Urak, Old Urak, Kahan, Chotair, Pasni Rek,
Gunduk, Bolan, Giridik; 1 & type Quetta) (all BM).

Distribution. — Only known from Afghanistan and W. Pakistan.

Subspecific characters. — g 11.7—13.5 (—14) mm. Male genitalia (Fig. 4, 5):
apex of cucullus much more pointed than in ssp. struvei, overlapping a great part of
the costa; dorsal surface of costa densely clothed with small spines, but coarser than in
ssp. struvei; ventro-distal end of costa unspined.

It is interesting to note that Reverdin, who, after Rambur, was the first to put the
chaos of the Hesperiidae in order, perhaps never saw a true geron. His sketch (1914) of
the valva of geron clearly shows an osthelderi, a species that was described eighteen years
later. Perhaps the same applies to Warren (1926, PI. 6 Fig. 5), but his photograph is
to dark to be decisive.

Spialia geron struvei Piingeler

Püngeler, 1914. — Iris 28 : 37, pl. 2 fig. 13, 20.
Type-locality: Barkul.

R. DE JONG: Systematics and evolution of Spialia 239

Distribution. — The few known localities of this species outside Afghanistan and
Baluchistan (see the distribution of the species).

Subspecific characters. — g' 13.5—14.2 mm. Male genitalia (Fig. 3): apex of
cucullus not strongly developed, just reaching the costa; dorsal surface of costa densely
clothed with very fine spines; ventro-distal end of costa spined.

Spialia doris (Walker)

Distribution (Map 4). — Widely but apparently highly discontinuously distributed
from Morocco to India.

In Morocco only captured in the following localities: El Aioun du Dràa (Rungs, 1945)
and High Atlas, Ziz Valley (Evans, 1949) and Ksar-es-Souk (material examined).
Several localities in N. Egypt, south to Heluan (Graves, 1925; Hemming, 1932b; Evans,
1949). From Jordan only known from Qasr Azraq and Hazin (Hemming, 1932b).
Widely distributed in W. and SW. Arabia, from Jidda southward, and in Africa from
Port Sudan and Kordofan to Somaliland (material examined) and N. Kenya (Baringo
District; specimen in National Museum, Nairobi, according to personal communication
by Mr. J. H. Lourens). SW. and S. Iran (Ahwaz, Kerman), Sind, Cutch, Punjab and
Rajputana (Evans, 1949; material examined). Hemming (1932b) recorded a male from
River Dyala (Iraq), but this specimen (in the BM) belongs to Spialia osthelderi.

Habitat. — Desert, as far as the food plant grows. In Yemen up to 2800 m (material
examined).

Biology. — In Egypt possibly two generations per year. The material in the BM dates
from the periods March—April and September—October. Reverdin (1914), however,
used for his description of amenophis (see below), a male caught August 1st and a male
caught November 11th (both from Heliopolis). The spring and autumn forms differ

= ZN

=f
g

Map 4 - Spialia doris

240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

in size.

In Arabia the species is known from all months in the period September to May, but
in summer the species seems to be absent, apart from a few specimens taken in July.
No size difference between the spring and autumn specimens could be established.

From other parts of the range the data are too scanty to allow for reliable conclusions.

The only known foodplant is Convolvulus lanatus (Graves, 1925: Egypt), a species
of the desert.

Geographic variation. — There is a geographic variation in the size, the colour of the
underside and the extension of the spots of the upperside.

Spialia doris daphne Evans

Evans, 1949. — Cat. Hesp. Eur. Asia & Austr. : 177.
Type-locality: Ziz Valley (High Atlas).

Material examined. — 13 & 5 9 :1 oO Ziz Valley, 21.1V.1935 (holotype) (BM),
12 g 5 9 Ksar-es-Souk, 8—18.V.1950, 3500 ft (CW, ML).

Distribution. — Only known from S. Morocco. Rungs (1945) recorded the species
from El Aioun du Drâa (S. Morocco) under the name “Pyrgus doris adenensis Butl.”.
This may be daphne, as this subspecies shows the greatest resemblance to subspecies doris
(adenensis is a synonym of doris).

Subspecific characters. — The original description reads: "Unh very dark greenish
brown, markings sharply defined, basal cell spot elongate. Upf basal cell spot elongate,
no spots in spaces 4 and 5: outer discal spot in space 1b vestigial. gt F 1214 mm.” This
description of the type is correct, except that the length of the fore wing is only 11.5 mm.
The other specimens, however, show some variation, viz. on the fore wing upperside
the discal spots in spaces 4 and 5 and the outer discal spot in space 1b may be present,
and the length of the fore wing varies from 11 to 12 mm.

Note. — It is a bad usage to describe subspecies on account of a single specimen.
Although it is to be expected that in a geographically isolated population differentiating
characters will develop, this is not necessary. As to the present instance, the type of ssp.
daphne could have been an extreme variation of a population that is otherwise similar to
ssp. doris, the more so as the variation concerns characters that are liable to variation
in ssp. doris. Fortunately, the distinction of ssp. daphne is not superfluous, but Evans
could not know that on account of his single specimen!

Spialia doris amenophis (Reverdin)

Reverdin, 1914. — Bull. Soc. Lép. Genève 3: 55—65, pl. 3 fig. 5, 11, pl. 4 fig. 1.
Type-locality: Heliopolis (Egypt).

Material examined. — 134 12 2. Spring form: 7 & 5 © Mokattam Hills (near
Cairo) (BM), 1 g 19 Heliopolis (BM, MW). Autumn form: 4 & 5 9 Qassasin,
1 9 Wadi Rished (near Heluan), 1 & Wadi el Tih (all BM).

Distribution. — Northern Egypt, from northern Sinai (Mehemdia; Graves, 1925) to
Heluan; Jordan (Qasr Azraq, Hazim; Hemming, 1932b).

Subspecific characters. — Spring form larger than autumn form: g' 11.1—12.4 mm

R. DE JONG: Systematics and evolution of Spialia 241

(9 up to 13.7 mm!) and g° 10.2—11.2 mm, respectively. On the upperside the
markings very broad. The discal spots in spaces 3 to 8 may be fused into an irregular
band, while in other forms the discal spots in spaces 4 and 5 are minute or absent.

Spialia doris doris (Walker)

Walker, 1870. — Entomologist 5 : 56.
Type-locality: Tajora, Red Sea.

Material examined. — 58 4 40 9: 42 g' 35 © Arabia (Jidda, Tihama Sabata,
Yemen, Aden, Hadramaut) (BM, MW), 4 & Perim, 2 g' French Somaliland (1 g°
Djibouti; 1 & holotype, Tajora), 6 g 3 $ Somaliland (Ghibdo River, Djibouti, Berbe-
ra, Hargeisa, Sheikh, Saleh Spring, Bihendula, Buran), 2 & 1 9 Ethiopia (Massowah,
Meiso, Dire Daoua) (all BM), 2 g 1 © Sudan (Port Sudan; Kordofan: Jebel Angageh,
Angageh Wells) (BM, MW).

Distribution. — The range of the species south of Jidda and Port Sudan: W., SW. and
S. Arabia, Sudan, Ethiopia, Somaliland.

Subspecific characters. — A rather constant form. On the upperside well-marked, but
all markings reduced as compared with ssp. amenophis, except the discal spots on the
upperside of the hind wing. The discal spots in spaces 4 and 5 on the fore wing upper-
side minute or absent. Submarginal spots may be faint. Ground colour of hind wing
underside dark greyish olivaceous, somewhat like ssp. amenophis, but sometimes more
yellowish. & (9—)10—11.6 mm.

Note. — Evidently overlooking the description by Walker, Butler (1884) described
the same form under the name Pyrgus evanidus var. adenensis from Aden (holotype in
the BM examined).

Spialia doris evanida (Butler)

Butler, 1880. — Ann. Mag. Nat. Hist. 5 (5) : 223.
Type-locality: Hubb River (S. Baluchistan).

Material examined. — 17 & 5 9 : 1g 1 9 S. Iran (Ahwaz, Kerman) (BM),
13 g' 3 2 Sind (Hubb River, type; Karachi, Hidrabad) (BM, MW, ML), 14 Cutch,
2 d Punjab (Campbellpore), 1 Q Rajputana (Deesa) (all BM).

Distribution. — The eastern part of the range of the species, from S. Iran eastward.

Subspecific characters. — Above like ssp. doris, below like ssp. doris but ground colour
paler, particularly in the hind wing which is pale yellowish. There exists, however, some
individual variation, while some specimens are as dark as ssp. doris. Smaller than the
latter, & 8.6—10.2 mm.

Note. — Of the four subspecies recognized here, doris and evanida are most alike.
Evans (1949) even considered the difference between them too small for subspecific
recognition. However, the populations of ssp. doris and ssp. evanida appear to be entirely
isolated geographically and I consider the subspecific distinction of evanida justified.

4. THE sertorius SPECIES GROUP

In the Palaearctic region the sertorius species group is represented by seven allopatric
forms, externally clearly distinguishable, but the differences in the genitalia are almost

242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

imperceptible or absent. These forms are: sertorius, ali, therapne, orbifer, hilaris, lugens
and carnea. Whether one considers these forms subspecies or separate species (perhaps
united into a superspecies) seems to be only a matter of taste, apart from the following
considerations.

(1). therapne is the forma of Corsica and Sardinia. Kauffmann (1955) recorded the
capture of sertorius at Evisa (Corsica). According to Bretherton (1966) sertorius was
recently found in two places in Corsica, and Higgins & Riley (1970) state that
specimens taken at Corte and Evisa have been described as intermediates of therapne and
sertorius.

(2). The second generation of the North African al? looks very much like therapne,
the difference being mainly the shape of the central spot and the colour on the under-
side of the hind wing.

(3). In sertorius, ali and therapne the shape of the central spot of the hind wing
underside is angular; in orbifer, hilaris, lugens and carnea this spot is rounded.

(4). In Central and Eastern Europe the forms sertorius and orbifer must approach
each other very closely and possibly overlap to a small extent. In Poland only sertorius
is known (Bleszynski e.a., 1965; Krzywicki, 1970). The only orbifer specimens examined
from Czechoslovakia originate from Jung-Bunzlau (Bohemia), further I saw only
sertorius from Czechoslovakia. According to Moucha & Novak (1960) only two reliable
records of orbifer are known from Slovakia (Stürovo, Koväcov), both near Hungary,
while all other specimens collected in Slovakia belong to sertorius (see also Reiprich,
1960). The easternmost locality from where I have seen sertorius is Szobrancz, near the
Czechoslovakian-Russian frontier.

Alberti (1965) said to have seen specimens with transitional characters from Northern
Hungary in the Hungarian Natural History Museum, but the specimens of both sertorius
and orbifer from the collections of this museum examined by me did not allow for such
a conclusion. The boundary between sertorius and orbifer in Hungary appears to be the
river Raba: all specimens collected west of it are sertorius (like all Austrian specimens),
while east of the river only orbifer is known to occur.

In Northern Yugoslavia sertorius is widely, but locally distributed. I have seen
specimens from the Sneznik (30 km north of Rijeka) and from Fuzine (15 km east of
Rijeka). Prof. Lorkovië kindly informed me that sertorius has been captured at some
more localities in northern Yugoslavia, viz. Zagreb and surroundings (Podsused, Samobor,
Japetië), Klek near Ogulin, Hrvatsko (see also Lorkovië & Mladinov, 1971), Slunj,
Ljeskovac (Plitvice lakes), Velika near Pozega, and Banja Luka. In the coastal region
sertorius and orbifer appear to approach each other very closely. At Trieste only sertorius
has been captured. Verity (1940) stated that orbifer occurs in Istria, but he did not give
exact localities and the reliability of this record is doubtful. The northernmost locality
in the coastal region from where I have seen orbifer is Senj (= Zengg). Koca (1901)
recorded the same form from the nearby Krizpolje. Lorkovië (in litt.) found orbifer at
Plitvice. Further south only orbifer has been collected (see also Lorkovië, 1973).

Rebel (1895) recorded specimens of the spring brood at Plitvice, with the spots on
the underside of the hind wing of the sertorius type, but with the ground colour olive
brown. It is not clear whether Rebel thought these specimens to belong to sertorius or to
orbifer. Very probably they do not indicate a hybrid population. Lorkovië (in litt.) found
several orbifer specimens at Plitvice and only one sertorius specimen, apparently without
intermediate characters.

R. DE JONG: Systematics and evolution of Spialia 243

Thus, in spite of the close geographic vicinity of sertorius and orbifer the occurrence
of intermediate populations has not yet been proved. According to the material available
at present a possible hybridization is a rare event. This idea is supported by the discovery
of differences between sertorius and orbifer in the larval and pupal stages in N. Yugo-
slavia (Lorkovië, 1973).

Exact localities in the border region of sertorius and orbifer from where I have seen
specimens or reliable records have been represented in Map 5.

(5). Throughout Peninsular Italy only sertorius is found, but in Sicily it is replaced
by an orbifer population with some sertorius features in the female (see under Spialia
orbifer).

(6). The forms hilaris and orbifer and the forms /wgens and carnea are connected
by intermediates in SE. Turkey and the Samarkand region, respectively.

Of course, these arguments are not conclusive, but from the distribution and dif-
ferentiation of sertorius, ali and therapne on one side and of orbifer, hilaris, lugens and
carnea on the other, one may conclude that these forms fall taxonomically and geo-
graphically into two groups, viz. a western serforius group and an eastern orbifer group.

Fig. 21—23. Female genitalia of Spialia sertorius sertorius (Chiclana, S. Spain). 21, ventral side;
22, left side; 23 right papilla analis

244 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

For practical reasons I treat the two groups as species and I am uniting them into the
superspecies sertorius. However, I must stress the fact that one could equally well consider
sertorius and orbifer subspecies groups of one species. For the evolutionary and geo-
graphic history of superspecies sertorius, that shows the arrangement adopted to reflect
the phylogenetic relationships, see Chapter 5.

In the Palaearctic no other representatives of the sertorius group exist. It is, however,
surprising that there are two species outside that region which are apparently closely
related to sertorius and orbifer and which show magnificently how the present type of
valva of sertorius and orbifer has developed. One of these species viz. mafa, is African,
and extends from South Africa to Abyssinia (Evans, 1937), SW. Arabia (Gabriel, 1954)
and the vicinity of Mecca (material examined). The other species, galba, occurs
throughout India, Ceylon and a part of Burma, and is also known from Hainan (Evans,
1949, 1956). Their valvae and gnathos are shown in Fig. 32—35. For the phylogenetic
relations between galba, mafa and superspecies sertorius, see Chapter 5. From a broad
point of view one could consider this species group a superspecies as the composing
species are allopatric. However, by accepting a superspecies sertorius with the species
sertorius and orbifer, it is impractical and obscuring history, to unite these species with
mafa and galba into a single superspecies.

SUPERSPECIES SPIALIA SERTORIUS (Hoffmansegg)

Taxonomy.

It is not difficult to distinguish the species sertorius and orbifer with external
characters. In sertorius the ground colour of the underside of the hind wing is red in
various shades, from ochreous to a very vivid red, and the spots on the underside of the
hind wing, particularly the central spot and the costal spot (in spaces 7—8) are more
or less angular. In orbifer the ground colour of the underside of the hind wing is
greenish or yellowish olive-grey, the spots are rounded and the costal spot is particularly
distinct. Only in the subspecies carzea, orbifer can have a reddish underside of the hind
wing, but as this subspecies is Asiatic it cannot be confused with sertorius.

In the male genital armature the difference between sertorizs and orbifer is much less
distinct. Moreover, the differential characters are subject to variation and not entirely
reliable. The differences were discovered by Warren (1926), who emphasized their in-
constancy. Kauffmann (1955) considered them too unreliable to be of any use, but this
is slightly exaggerated.

The differences can be found in:

(a). Gnathos (Fig. 25, 27, 29, 31). The ventral spined part is narrowly pointed in
sertorius ali, less narrowly pointed in orbifer, bluntly rounded in sertorius therapne and
rounded in sertorius sertorius. Especially in s. sertorius the variation is considerable and
specimens with a therapne or orbifer-like gnathos are not extremely rare. Further, the
left and right parts of the gnathos may differ in form. Moreover, the angle from which
the structure is observed, is important, an oblique position of the gnathos causing a more
pointed appearance.

(b). Hairy fold at the inside of the valve. (Fig. 24, 26, 28, 30). In orbifer the
ventral, horizontal part is usually best developed, in sertorius the horizontal and vertical
parts are about equally well developed or the vertical part is best developed (sertorius

R. DE JONG: Systematics and evolution of Spialia 245

Fig. 24— 29. Inside of right valva and outside of left part of gnathos of Spialia. 24—25, S.
sertorius sertorius (Hannover); 26—27, S. sertorius ali (Algeria); 28—29, S. sertorius therapne
(Corsica)

ali). The variation in this character does not depend on the angle from which the fold
is observed.

In the female genitalia of sertorius and orbifer (Fig. 21—23) I could not find clear
differences.

From these facts it is obvious, that a possible hybrid between sertorius and orbifer
cannot be detected by the genital characters, and only by the external ones.

246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

a 2 == yA \

(SSS. ; Er N
SQW
xè SS

Fig. 30—35. Inside of right valva and outside of left part of gnathos of Spialia. 30—31, S. orbifer
orbifer (Turkey); 32—33, S. mafa (S. Africa); 34—35, S. galba (Ceylon)

Spialia sertorius (Hoffmansegg)

Nomenclature. — Apart from sertorius, two names have been used for this species,
viz. sao Hübner and hibiscae Hübner. Warren (1926) pointed out that Papilio sao
Hübner, 1800— 1803, is a junior homonym of Papilio sao Bergstrasser, 1779, and that
sertorius Hoffmansegg, 1804, is the correct name. However, Hemming (1936) recorded
the name hibiscae, attributed by him to Hübner (1790—93). This name has been used in
important publications, such as Verity (1940, 1947) and Kauffmann (1951). Evans

R. DE JONG: Systematics and evolution of Spialia 247

(1947) indicated that there is no evidence that the paper “Lepidoptera Linnei” where
Hemming found the name hibiscae, had ever been published, so that the name hibzscae
Hemming, 1936, should be placed as a synonym of sertorius Hoffmansegg, 1804.

Some authors (e.g. Lempke, 1953) have made an efffort to alter the gender of the
name sertorius in accordance with that of Spialia, resulting in the combination Spialza
sertoria. This is incorrect, sertorius not being an adjective but a proper name.

Distribution (Map 6). — NW. Africa; from S. Europe northward to southern
Bretagne (Picard, 1950), southern Netherlands, Osnabrück, Hannover and Harz
Mountains, east to southern Poland (Krzywicki, 1970), the Czechoslovakian— Russian
frontier, the river Raba in Hungary and Slavonia and Croatia in N. Yugoslavia. For
details of the eastern distributional limits, see above and Map 5. Spialia sertorius occurs
throughout Italy, Corsica and Sardinia, but in Sicily it is replaced by orbifer.

Habitat. — Flowery slopes, dry meadows from the lowlands up to 2200 m in the
French Pyrenees Rondou, 1932), to 2400 m in Spanish mountains (Manley & Allcard,
1970) and to 2300 m in Switzerland (Schmidlin, 1949). Mainly in xerotherm limestone
environments, undoubtedly in relation with the requirements of the food plants.

Biology. — Everywhere two generations a year, flying from April to June and from
July to September, but in mountainous districts often monovoltine. The larva hibernates
(Forster & Wohlfahrt, 1955).

Food plants: Sanguisorba minor, Potentilla verna, Rubus idaeus (Rosaceae) (Verity,
1940; Forster & Wohlfahrt, 1955). The food plants of the subspecies alt and therapne
are unknown.

Geographic variation. — Three subspecies are recognized here.

Spialia sertorius ali (Oberthür)

Oberthiir, 1881. — Etude d’Ent. 1 : 61, pl. 2 fig. 3.
Type-locality: Lambèse (Algeria).

Material examined. — 231 g' 114 9 : 38 4 17 @ Morocco (Tanger, Tetuan, Azrou,
Ketama, Timhadit, Ifrane, Ras-el-Ma, Valley of R’dat, Tamarouth, Asni, Ain Leuh,
Mrassine, Marchand) (BM, ML), 193 g 96 @ Algeria (Batna, Djebel-Aurès, Lambese,
Souk-Ahras, Tlemcen, Géryville, Sebdou, Hammam R'irha, Oran, Titen Yaya, El Hajeb,
El Kantara, Djelfa, Khenchela, Algiers, Tafna, Aflou, Blido, El-Maouna, Nedroma, El
Qutaya, Lafla-Marnia, El-biar, Hussein Dey, Sidi-bel-Abbès, Duffana) (BM, ML), 1 9
Tunisia (Ain-Draham) (BM).

Distribution. — Confined to NW. Africa.

Subspecific characters. — Male genitalia: gnathos ventrally narrowly pointed. External
characters: first brood, & 8.4—11.6 mm, underside of the hind wing red or pale brown
with dark striae, at the termen alternating white and red or brown streaks, because the
submarginal spots are modified, central spot with long projections towards base and
termen; second brood: g' 8.3—10.3 mm, underside of the hind wing mostly yellowish
red with less pronounced white markings, on the upperside of both wings the wite
markings with a yellow shade.

Note. — The second brood is very similar to subspecies therapne and for this reason
Oberthiir (1910) gave it the name therapnoides. It can, however, easily be separated by
the less warm ochreous red underside and by the central spot of the underside of the

248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

hind wing, that in therapnoides has long projections towards base and termen (as in the
first brood), while in therapne it only has a projection towards the termen.

Spialia sertorius therapne (Rambur)

Rambur, 1832. — Ann. Soc. Ent. France 1832 : pl. 7 fig. 4.
Type-locality: Corsica.

Material examined. — 54 g' 21 9 : 52 g 20 Q Corsica (BM, ML, HO, HC),
2 01 ® Sardinia (BM, ML).

Distribution. — Confined to Corsica and Sardinia.

Subspecific characters. — Male genitalia: ventral, spined part of gnathos not as narrow
as in ssp. ali, blunt but not rounded as in sertorius. External characters: very small,
d' 7.8—9.9 mm; on the upperside shaded yellow, spots yellowish, submarginal spots very
faint or absent. On the underside warm ochreous red, spots arranged as in ssp. sertorius;
see also ssp. ali. i

Note. — According to Higgins & Riley (1970) therapne is bivoltine, flying in April
and September. However, I have only seen specimens dated from June to September.

Spialia sertorius sertorius (Hoffmansegg)

Hoffmansegg, 1804. — Mag. f. Insektenkunde (Illiger) 3 : 203.
Type-locality: Germany.

Material examined. — 779 G 420 ® : 36 g' 13 9 Portugal (BM, ML), 49 &
14 9 S. Spain (BM, ML, ITZ), 101 & 33 9 C. and N. Spain (BM, ML), 10 g
3 Q Spanish Pyrenees (BM, ML), 92 & 45 9 French Pyrenees (Hautes Pyr., Pyr. Or.)
(BM, ML, ITZ), 231 & 148 Q France (BM, ML, ITZ), 13 & 8 2 Belgium (BM,
ML), 12 g 10 2 Netherlands (ML, ITZ), 44 & 16 9 Switzerland (BM, ML, ITZ),
34 g' 14 Q Germany (Bayern to Hannover and Leipzig) (BM, MB, ML, ITZ), 39 g
35 Q Austria (BM, ML, MB), 7 g' 8 @ Czechoslovakia (Praha, Slany, Hlubocepy,
Trencsén, Eperjes, Szobrincz) (MB, ML, BM), 1 g 1 © Hungary (Sopron, Magyarovär)
(MB), 12 g' 4 9 N. Italy (BM, ML, ITZ), 84 g° 63 9 C. Italy (BM, ML), 1 2
Elba (ML), 2 g 1 @ Calabria (BM), 10 g° 2 2 Trieste and Istria (BM, ML, MB),
1 g' Slovenia (Sneznik) (ML), 1 4 1 2 Croatia (Fuzine) (ML).

Distribution. — The whole distribution area of the species, except NW. Africa and
Corsica-Sardinia, i.e. West, Central and South Europe, east to Slavonia and E. Czecho-
slovakia; for details, see above and Map 5.

Subspecific characters. — Male genitalia: ventral, spined end of gnathos broad and
rounded. There is some variation in the direction of ssp. ali and ssp. therapne, but these
forms are rarely equalled. External characters: on the underside of the hind wing the
submarginal spots not modified as in ssp. 4/7, ground colour red in various shades to
ochreous red or, less commonly, yellowish, particularly in the second brood in S. Europe.
The two broods differ in size: first brood 4 10—12 mm. second brood g' 9—11 mm,
but there is much variation and a general rule cannot be given.

Variation. — This subspecies is rather variable, especially in Spain. Some authors,
e.g. Verity (1940), have recognized several subspecies or “races”. At least partly, this

R. DE JONG: Systematics and evolution of Spialia 249

Map 5. Distribution of Spialia sertorius and orbifer in Central and Southeastern Europe. @ =
sertorius, material examined; O = sertorius, literature records; À = orbifer, material examined;
A = orbifer, literature records.

250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

variation does not seem to be geographic, but it is possible that some geographic variation
exists. The following variation is known:

(1). The spots on the upperside may be all present and well-marked or may be
reduced. Particularly the submarginal spots can be totally absent. Based on this variation
Verity (1940, 1947) and Kauffmann (1951) recognized the subspecies hibiscae (by
Verity attributed to Hübner, by Kauffmann to Hemming; see also above, under Nomen-
clature) (well-marked) and sertorius (markings reduced). The second brood of both
forms is somewhat smaller than the first brood and has been named parvula Verity (for
sertorins) and minor Rebel (for hibiscae).

Both Verity and Kauffmann state that the dark form (sertorius) is confined to humid
places; therefore, it can hardly be assigned subspecific state. There are still more reasons
to mistrust the correctness of the subspecific separation. In most places both forms occur
together, connected by many transitions. I have seen both forms from Lisbon, the
Pyrenees, the Basses Alpes, Belgium, the Netherlands, Germany and Lower Austria.
Mostly hibiscae is the dominant form, but sertorivs can dominate in some localities (and
perhaps only in some years). Bergmann (1952) found proportionally more hibiscae when
the years were drier.

Lempke (1953) is correct in stating that in the Netherlands the dark form dominates,
but he also mentions better marked specimens and says (p. 244): “Such specimens agree
totally with subsp. hibiscae Hemming (...). It is difficult to name such specimens as,
with us, they belong to the infrasubspecific category, while the name was given to a
group of higher rank. Nomenclaturally these limits are, of course, splendid, but in
practice all limits disappear” However, in this case as well as in many others the dif-
ficulties arise from a wrong interpretation of the facts and a wrong concept of super-
species. In the present case, sertorius and hibiscae are not separate subspecies, but climatic
forms. Consequently, I do not recognize hibiscae as a separate subspecies.

(2). Mostly, the ground colour on the underside of the hind wing is red or reddish,
but in southern populations the colour may be ochreous yellow or even pale yellow. A
yellow colour can also occur in the second brood in northern populations. The cause of
this variation is unknown. As the yellow colour is not confined to any particular region,
it cannot be used as a subspecific character.

(3). Particularly in southern regions there is a great variation in size, which is partly
individual, partly geographic. In Spain larger and smaller specimens fly together (Warren,
1926). According to Zerny (1927), three size categories can be distinguished in the
vicinity of Albarracin. This suggests that no transitional specimens occur, but that is not
true. The difference can be very large: four males from San Ildefonso (Segovia, June to
August) measured 9.8, 10.1, 12.2 and 13.1 mm, four females from the same locality
10.2, 13.1, 13.4 and 14.2 mm (BM).

In Peninsular Italy, except S. Calabria, all specimens are small, first brood & 9.711
mm, second brood g' 8.4—10.1 mm. In N. Italy and S. Calabria the specimens measure
as much as 12—13 mm.

(4). Some minor variation exists as mentioned below. Apart from sertorius and
hibiscae the following forms have been described as “races”:

(a). gracilis Verity, 1921 (type-locality: Florence). The small form of Peninsular
Italy; on the upperside spots small but all present; underside of the hind wing red or pale
yellow, with relatively few transitional specimens. The first generation (for measurements
see above, under (3)) was named szbgracilis by Verity (1921). This form is rather

R. DE JONG: Systematics and evolution of Spialia 251

constant in Italy, although the larger specimens of the first brood cannot be distinguished
from hibiscae. In Spain gracilis occurs together with larger specimens, in France it occurs
only in dry places in the south. Small specimens with pale underside of the hind wing
were already known under the name eucrate Ochsenheimer, 1805. Such specimens can
also be found in the second brood in northern populations.

At the moment I do not know how to place the form gracilis. As it does not seem
to have a distributional area of its own, it appears inappropriate to give it subspecific
rank. Nevertheless, it is not impossible that gracilis originated by geographic isolation
(see also Chapter 5).

Note. — Verity described gracilis already in 1919 as the second generation near
Florence. According to the International Code of Zoological Nomenclature, Article 45d,
the original status of gracilis is determined as infrasubspecific. However, in 1921 Verity
clearly applied the name to a particular geographic area and he called gracilis a “race”.
Therefore, the subspecific status of the name dates from 1921.

(b). gwadarramensis Warren, 1925 (type-locality: La Granja, Sierra de Guadarrama).
The largest form known, according to Warren (1925) 30—31 mm (centre of the thorax
to tip of the fore wing x2). I saw only one female from the type-locality, that measured
14.2 mm. Furthermore, the form is characterized by the ground colour of the underside
of the hind wing, which is vivid red, much deeper in shade and more brilliant than in
Central European specimens. This, too, is the case in the single female seen from La
Granja. However, as stated by Warren, gwadarramensis occurs together with normal-sized
and small specimens. I saw two males from Sierra de Guadarrama that were almost of the
size of gracilis. Moreover, not all large specimens from Central Spain are richly coloured,
e.g. the large male and female from San Ildefonso mentioned above (under (3)) have
the ground colour of the hind wing underside dark yellow and brick red, respectively.

Like gracilis, this variety cannot be defined as a subspecies.

(c). gavarniensis Warren, 1926 (type-locality: Gavarnie). According to Warren,
almost all specimens in the vicinity of Gavarnie (Hautes Pyrénées) differ from ssp.
sertorius in the warm yellow-brown, almost orange colour of all parts of the underside,
which are white in ssp. sertorius with the exception of the white spots. The ground colour
of the underside of the hind wing of gavarniensis is brighter red than of sertorizs.

Indeed, most specimens from Hautes Pyrénées show this type of variation, but rarely
in an extreme form and many specimens are practically indistinguishable from Central
European sertorius.

I have seen a female from Ordesa, at the Spanish side of Cirque de Gavarnie, that
looked like gavarniensis, but another female and two males from the same locality did not
show the colour that is typical for gavarniensis.

The cause of this variation which is confined to the Central Pyrenees, is unknown.
It is not very likely that it originated during geographic isolation, as at least at present
sertorius is wide-spread in the Central Pyrenees, flying from the lowlands up to 2200 m.
At any rate it originated postglacially, as the Pyrenees were uninhabitable for sertorius
during glacial periods. For this reason and for the fact that in the context of the total
variation of the species gavarniensis is only an unimportant minor variety, it is undesirable
to give it the same rank as the forms ali and therapne. I am placing it here as a local
form.

(d). alioides Verity, 1926 (type-locality: Oulx, Cottian Alps). Central spot on the
underside of the hind wing large with projections towards base and termen. Ground

252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

colour of the underside of the hind wing ochreous or yellowish, whitish along termen,
veins outlined in a paler shade. Upperside as ssp. sertorius, but the specimens are larger,
up to 13 mm. Such specimens occur in the western Alps, but mostly the colour of the
underside of the hind wing is red or ochreous red and all transitions to sertorius occur.
This form seems to be still less sharply defined than gavarniensis and I regard it also a
local form.

According to Verity (1940) the same form occurs in S. Calabria. I have seen only
2 4 and 1 9 from S. Calabria and these specimens looked like sertorius with some
variation towards the Sicilian orbifer.

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of 4 ||] | | (ll ID
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H Map 6
© 0 “ sertorius species group
EH Spialia sertorius
Ÿ Im Spialia orbifer

SS Sp'alia mafa
Za Spialia galba

Spialia orbifer (Hübner)

Distribution (Map 6). — Sicily; from N. Yugoslavia, Hungary and Czechoslovakia
eastward through S. Russia and S. Siberia to the Amur region and N. Thibet (Sining)
and through the Balkans and Turkey to Baluchistan and Afghanistan from where it goes
north through Turkestan to S. Siberia.

The distribution limits are badly known. For the distribution in E. Europe, see above
and Map 5. Although it is very probable that this distribution is continuous, there are no
records from large intervening areas in Russia and Siberia. From Russia we have the
observation of Caradja (1895), that orbifer is widely distributed in SW. Russia and. the
Volga region, further the record of Eversmann (1844), who mentioned it from Saratov
on the Volga. Krulikowsky (1908) recorded orbifer from Kasan and Spassk, but he only
relied on some literature records. The species is known from the Crimea (Melioransky,
1897; Korshunov, 1964; material examined), the Caucasus and Transcaucasia (Romanoff,
1884; Alberti, 1969), but in the S. and SE. Russian steppe region it is apparently rare,
as it is not mentioned, e.g., by Obraztsov (1936) from the Transdnjepr region, neither
by Alberti & Soffner (1962) from S. Russia, nor by Gross (1925) from Chwalynsk
on the Volga.

From the W. Siberian lowlands it is only recorded by Sjtsjuko (1916) from Tjumen.

R. DE JONG: Systematics and evolution of Spialia 253

Further east it is known from the upper Irtysch (Lederer, 1853), Altai and Kentei
(Elwes, 1899; material examined) and it is widely distributed though uncommon in the
Amur region and the Maritime Province (Staudinger, 1892; Kurentzov, 1949, 1970).
Matsumura (1927) and Bryk (1946) did not record orbifer from Korea, but Sugitani
(1936, see Evans, 1949) described the form murasaki from that country. In China it is
undoubtedly much more widely distributed than is suggested by the few localities known
(Shansi, S. Shensi, Sining; see material examined).

In the southern Palaearctic orbifer is widely distributed. The southernmost localities
are: vicinity of Jerusalem (Graves, 1925; material examined), Bished (Iran) (Forster,
1939) and Ziarat (W. Pakistan) (material examined). Through Afghanistan, Trans-
caspia and Turkestan to the Ili region (Alphéraky, 1881; Wagner, 1913) and possibly
continuously to the Altai. For details, see below.

Habitat. — Like the habitat of Spzalia sertorius. In Macedonia I found orbifer up to
2000 m, in C. Anatolia it flies up to 1400 m (Pfeiffer, 1927), in Chitral it is not rare
at 4000 m (Evans, 1927).

Biology. — Bivoltine in the western part of the area, west of Iran. According to
Pfeiffer (1932) a third brood occurs near Marash (E. Turkey), but Graves (1925) is of
the opinion that late specimens may rather be of the second brood, but delayed by the
summer pause. Evans (1949) assumed that the Asiatic populations (from Transcaspia
eastward) are single-brooded.

The larval stages remained unknown till, very recently, Lorkovié (1973) succeeded in
rearing this species on Sanguisorba minor (Rosaceae).

Geographic variation. — Apart from the variation in number of broods per year, the
species varies in size, colour and markings. The variation in C. and E. Asia is still badly
known and with more material available subspecies lugens may prove to be heterogeneous.

Spialia orbifer orbifer (Hübner)

Hiibner, 1823. — Samml. Eur. Schmett. 1, pl. 161 fig. 803—806.
Type-locality (designated here) : Hungary.

Material examined. — 424 G 187 9 :10 & 6 @ Sicily (Messina, Castelbuono in
Madonie, Palermo, Etna-Randazzo) (BM), 1 & “Istria” (ML), 55 & 27 ® Hungary
(Oski, Fenyöfó, Gyenesdiás, Szár, Ugod, Pilisvörösvár, Sümeg, Akali, Budapest) (MB,
ML, BM), 1 & 1 2 Bohemia (Jung Bunzlau) (BM), 7 g 1 9 Croatia (Zengg= Senj)
(BM), 10 & 7 © Bosnia (BM, ML, ITZ), 1 g' Hercegovina (ML), 9 g' 7 9
Dalmatia (BM, ML, ITZ), 81 8 26 9 Yugoslavian Macedonia (ML, HC, ITZ),
102 4 60 9 Greece (BM, ML, ITZ), 1 g Crimea (BM), 143 8 48 © Turkey
(European and Asiatic) (BM, ML), 3 9 Russian Armenia (Vartian), 3 g 1 9 N. Iran
(N. of Teheran) (BM, Vartian).

Distribution. — The western part of the range, eastward to W. Siberia, south-eastward
to N. Iran (north of Teheran). Also in Sicily.
Subspecific characters. — Bivoltine. First brood, g' 11—12.6 mm, second brood,

d' 8.4— 11 mm. Markings on the upperside variable, sometimes all spots present, some-
times they are very reduced, particularly the submarginal spots. Ground colour on the
underside of the hind wing greenish to ochreous olivaceous.

Variation. — The name minor Rebel, 1909, for the second brood of Spialia sertorius,

254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

is also suitable for the small brood of orbifer. In a series of about 50 specimens collected
in Mavrovo (Yugoslavian Macedonia) in the first half of July, the males measure 10.5—
12.2 mm. As the specimens were collected between 1000 and 2000 m, it seems possible
that a single brood exists there (and perhaps in other mountain regions).

A dark, less marked form was described as tesselloides by Herrich-Schäffer (1845)
from Sicily. Verity (1940) gave some confusing remarks about the type-locality.
Evidently, he did not examine the original description by Herrich-Schäffer and relied
entirely on Keferstein (1851 : 326). The latter only mentioned that tesselloides occurs
in “Türkei”, without stating that it does not occur elsewhere. Herrich-Schäffer received
the type specimens from Keferstein and he clearly stated that they originated from Sicily.
Verity supposed that Herrich-Schäffer gave “S. Europe” as type-locality and that this was
corrected by Keferstein in “Turkey”. As this form occurs in Turkey, Saloniki, the
Olympus and Sicily (according to Verity), the problem is not very important.

The smaller second brood of tesselloides was named posttesselloides by Verity (1938)
after specimens from the Olympus. Evans (1949) correctly placed tesselloides as a name
relating to orbifer, but his opinion that posttesselloides is a synonym of sertorius is an
apparent mistake.

This dark form bears the same relation to the better marked nominate form, as the
form sertorius does to the well-marked form hibiscae. Also its valuation must be the same,
as both forms occur together over the main part of the distribution area, tesselloides
perhaps dominating in some districts, but without a distribution of its own and connected
with better marked specimens by innumerable transitions.

From the point of view of geographic history the occurrence of orbifer in Sicily is
highly interesting. The species seems to be scarce there. Verity (1940) saw only two
Sicilian specimens which he attributed to the form fesselloides. Indeed, all Sicilian males
in the BM are dark on the upperside, but the females are somewhat more distinctly
marked (this is a normal feature in orbifer). The hind wing underside of the males is
of the colour and markings of orbifer, but the central spot is rather angular. The females,
however, vary from ochreous to red and the central spot is very suggestive of sertorius.
The measurements are: g' 10.5—12.7 mm, 9 12.2—13.8 mm. In all, it seems advisable
to refer the specimens to orbifer, but they suggest a hybrid population rather than pure
orbifer.

Spialia orbifer hilaris (Staudinger)

Staudinger, 1901. — in Staudinger & Rebel — Cat. Lep. Pal. : 96.
Type-locality: Mardin (SE Turkey).

Material examined. — 115 g 41 Q : 80 8 34 ® Syria and Lebanon (Akbès,
Aleppo, Bscherre, Beirut, Damascus, Tripoli, Antiochia, Ain-Zahalta, Kassab, Bludan,
Deir Billa, Hasbaya, Zahlé) (BM, ML, ITZ), 16 g 5 9 Israél (BM, ITZ), 10 g
2 Q Iraq (Shaqlawa, Dohuk Mts., “Iraq”) (BM), 9 & W. Iran (Harir, Kermanshah,
Karind) (BM).

Distribution. — From SE. Turkey southward, to Matta (15 km SW. of Jerusalem;
material examined) and Amman (Jordan; Hemming, 1932b) and eastward to Kerman-
shah (W. Iran; material examined).

Subspecific characters. — Bivoltine. First brood G' 10.8—13.8 mm, second brood

R. DE JONG : Systematics and evolution of Spialia 255

d' 9—11.3 mm. On the upperside, the markings are more conspicuous than in ssp.
orbifer, particularly the submarginal spots.

Variation. — The small second brood was named secunda by Graves (1925). In NW.
Syria (Akbès) also less conspicuously spotted specimens occur which form a transition
to ssp. orbifer.

Note. — If compared in series ssp. hilaris is quite distinct from ssp. orbifer. Its
distribution, however, suggests, that it is a climatic form, adapted to hot and dry con-
ditions, rather than a subspecies (though climatic forms and subspecies are, of course, not
mutually exclusive). Provisionally I follow common use by considering hilaris a sub-
species. It would be interesting to have more material from the contact zone between the
subspecies orbifer and hilaris in order to study a possible clinal variation along a
temperature and/or drought gradient.

Spialia orbifer lugens (Staudinger)

Staudinger, 1886. — Stett. ent. Zeitung 47 : 256.
Type-locality: Ferghana.

Material examined. — 71 4 19 2:3 & NE. Iran (Khush Yailag) (WLB), 2 &
1 Q Transcaspia (Merv, Achal-Tekke) (BM), 1 g° Bokhara (BM), 8g 5 9
Samarkand (BM, ML), 1 & Hissar Mts. (ML), 5 & 2 9 Ferghana (Namangan,
Margelan, Gultscha) (BM), 24 & 4 $ Naryn (BM, ML), 9 g' 1 9 Talass Ala-Tau
(ML), 1 & Tashkent (ML), 1 g° Alexander Mts, 1 g 1 Q Issyk-kul, 2 g' Tian-
shan, 3 & Altai (Ongodai), 1 4 Amur region (Tjutju-ho), 5 g 4 ® Shansi, 3 &
S. Shensi, 1 & 1 © Sining (all BM).

Distribution. — The whole Asiatic range of the species east of the Caspian Sea,
except Afghanistan and from Baluchistan to Chitral.
Subspecific characters. — Monovoltine. Large, & and 9 to 14.6 mm, and dark on

the upperside, submarginal spots mostly faint, may be absent, but sometimes (mainly in
females) well-developed. On the underside of the hind wing indistinguishable from
ssp. orbifer.

Variation. — Size is variable, large specimens are found in China, g' 12.6—14.6 mm,
and in the Talass Ala-Tau, ¢ 13.3—14.1 mm, specimens from Naryn are somewhat
smaller, g 11—13.7 mm, the few males from the Altai measure only 11.9—12.6 mm.
Therefore, one can also refer the Altai specimens to ssp. orbifer, at least with regard
to their size.

Wagner (1913) remarked that orbifer occurred in the Ili region “sowohl in typischen
Stücken, als auch in der grösseren und dunkleren var. Lugens Stgr.” He also mentioned
specimens with large spots which he called “var. Hilaris Stgr.”. From these facts it seems
possible that the subspecies orbifer and lugens meet in N. Turkestan and the Altai.

In some females from the province of Samarkand, the ground colour of the underside
of the hind wing is reddish brown instead of greenish. Although such specimens have
the submarginal spot in spaces 4 and 5 at the underside of the hind wing well-developed
(unlike the usually less conspicuous spot in ssp. carnea), they are suggestive of ssp.
carnea. Possibly there is a large transitional zone between the subspecies lugens and
carnea. However, while I have seen ssp. lugens from Merv and the Achal-Tekke region,
ssp. carnea is recorded from the vicinity of Herat in W. Afghanistan (Clench &

256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

Shoumatoff, 1956).

Unfortunately, the material from N. Iran is too scarce to allow for a statement about
the contact zone between the subspecies orbifer and lugens. Three males from the
mountain Khush Yailaq in the eastern part of the Elburz Mountains are indistinguishable
from lugens specimens from Turkestan. Two males and one female from Derbend and
Vanak (N. of Teheran), about 400 km west of Khush Yailaq, look like orbifer
specimens from more western localities, without any trace of the influence of ssp. lugens.

Spialia orbifer carnea (Reverdin)

Reverdin, 1927. — in Bang-Haas - Horae Macrolep. 1 : 52, pl. 7 fig. 28.
Type-locality: Paghman Mountains (Afghanistan).

Material examined. — 36 & 14 9 :15 & 5 2 Baluchistan (Ziarat, Urak, Old Urak)
(BM), 4 & 3 9 Afghanistan (Paghman Mts, Arbarp, both W. of Kabul; Panjshir
Valley, N. of Kabul) (BM, ML), 14 @ 5 2 Chitral (Chitral, Shandur plateau) (BM,
CW), 3 & 1 2 Gilgit (Ghizar, Hailtar, Gilgit) (BM).

Distribution. — Afghanistan and from Baluchistan (vicinity of Quetta) to Chitral.

Subspecific characters. — As ssp. lugens, but on the underside of the hind wing warm
red to brown-yellow. g'° 12.2—14.2 mm, ® 13.6—16.4 (!) mm. In most specimens the
submarginal spot on the underside of the hind wing in spaces 4 and 5 is less conspicuous
than in other subspecies.

Variation. — On the upperside, the spots are variable, the submarginal spots may be
well-developed, but are sometimes totally absent (Chitral).

5. EVOLUTIONARY AND GEOGRAPHIC HISTORY OF THE GENUS Spialia
IN THE PALAEARCTIC REGION

a. General

In this chapter we will attempt to trace the evolutionary history of Spialia in the
Palaearctic by analyzing some characters as to their plesiomorphous (primitive) or
apomorphous (derived) condition (Hennig, 1966). The conclusions arrived at may serve
for the reconstruction of the geographic history. The reconstruction of the evolutionary
and geographic history forms the theoretical basis for the arrangement of the species
and subspecies adopted in the preceding chapters.

The phlomidis and sertorius groups will be dealt with separately. Their
relationship cannot be treated without a revision of the whole genus.

b. The phlomidis group

1. Evolutionary history.

We will at first examine the characters that are peculiar of the phlomidis group in
order to show that the arrangement adopted in this paper is phylogenetically correct.
Thereafter, we will try to detect the evolution within the phlomidis group.

(1). Valva. In the phlomidis group the cucullus is strongly spined at its apex. In
the rest of Spialia this character only occurs in spio, but less strongly developed. In the

R. DE JONG: Systematics and evolution of Spialia 257

other Spialia species the cucullus is quite different, more or less concave dorso-distally
and often forms a flap so as to envelop partly the costal process; spines as in the
phlomidis group are absent, but sometimes the rim of the flap or the extreme apex of the
cucullus is slightly indented. If we assume that the cucullus originally was a plain
structure, merely the ventral thickening of the valva, the phlomidis type with its bent
and spined apex is an apomorphous stage of the original, simple cucullus, just as the
type of cucullus with a flap is its apomorphous stage (but apparently developed at a later
time; this however, falls outside the scope of this paper).

Unless we assume that this character arose independently in the species of the
phlomidis group and in spio, the phlomidis group plus spio can be considered a mono-
phyletic group, as the species share the same apomorphous condition of the cucullus and
as they comprise all species with this condition.

(2). Aedeagus. Outside the phlomidis group a sclerotized excrescence of the aedeagus
is rare in Spialia. This part occurs in e.g. nanus and abscondita as a short, spined ventral
branch, quite different from the latero-ventral crest or branch (somewhat to the left),
found in the phlomidis group. Moreover, in the phlomidis group the distal end of the
aedeagus is bent downward to the left and strongly developed, a unique feature in Spialia.
Undoubtedly, an excrescence points to an apomorphous condition with regard to the
situation where this structure is absent. Evidently, it evolved in the phlomidis group and
in nanus and its allies along different lines. Also the type of apex found in the
phlomidis type of aedeagus must be the apomorphous condition and the simple, straight
aedeagus the plesiomorphous one.

So again we can state that the phlomidis group is monophyletic, but now spio falls
outside the group.

(3). Antevaginal structures. The large, wing-like structures of the phlomidis group
are unique in Spialia, but in spio a less developed type is found. The presence of such
specialized structures appears to be an apomorphous condition as compared with their
absence (3a) and the strong development in the phlomidis group appears to be an
apomorphous condition as compared with the weak development in spzo (3b).

Again, we find that the phlomidis group is monophyletic and that spzo must be
included, if we draw the limits more broadly.

Summarizing, it appears justified to call the species of the phlomidis group mono-
phyletic, originating from a common ancestor and comprising all progeny of that
ancestor known. Spialia spio is phylogenetically related to the phlomidis group, but it
does not share the apomorphous condition of some characters. For this reason and for its
distribution (Ethiopian against the eremian regions of the Palaearctic for the phlomidis
group) spio can better be left out of the phlomidis group.

Now, we will try to trace the evolution within the phlomidis group.

(4). Aedeagus. The excrescence of the aedeagus forms a crest in phlomidis, doris
and osthelderi and a branch in geron. We may assume that an outgrowth evolves from
small to large. Thus the situation in geron is the apomorphous condition and the crest,
the plesiomorphous one.

(5). Cucullus. The apex of the cucullus is scarcely bent in phlomidis but strongly
bent in the other species. Apparently, the larger the bend, the more advanced the
evolution. Thus the strongly bent condition appears to be apomorphous. =

(6). Costa of valva. The costa is dorsally more or less densely set with small spines
in osthelderi, geron and doris, smooth in phlomidis. These spines are unique in Spialia.

258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

They appear to represent the apomorphous condition with regard to the smooth dorsal
surface of the costa.

(7). Costa of valva. In doris the distal end of the costa is concave. This appears to
be a further differentiation of the costa that originally was only a thickening of the dorsal
ridge of the valva. Consequently, the condition in doris is apomorphous.

(8). Markings on the underside of the hind wing. The direction of the central band
on the underside of the hind wing is rather variable in Spialia, but usually the band is
connected with or directed to the outer spot in space 7 or to a point between the outer
and inner spots in space 7. In spio, osthelderi, geron and doris the central band is directed
to the outer spot in space 7 or somewhat more to the base, in phlomidis it is connected
with the inner spot in space 7. The lastnamed condition appears apomorphous, unless we
assume that the other condition arose independently in several species.

(9). Ground colour of the underside of the hind wing. In Spialia this ground colour

(Gp) =>} > O ® =)

da (av a es

A.) perg oO

ex Yn =

5 ae

LUI TEEN

10a

Fig. 36. Supposed evolution in the Spialia phlomidis species group. The numbers indicate characters
as indicated in the text. O = plesiomorphous condition, ® = apomorphous condition

R. DE JONG: Systematics and evolution of Spialia 259

is never white or whitish, except in osthelderi osthelderi. Evidently, the white colour is
the apomorphous stage. As mentioned above (chapter 3) the same character occurs in the
same region in some other Hesperiidae.

(10). Costa of the valva. In the phlomidis group the ventro-distal end of the costa
bears strong spines, except in geron geron. Generally, the absence of spines on the valves
in Spialia appears to be a less differentiated, plesiomorphous condition. In the case of
g. geron, however, we must suppose that the spines have been lost, for the following
reason. In Spialia usually the ventral or ventro-distal part of the costa is differentiated
with spines or brushes. In phlomidis, doris, osthelderi and geron struvei the position and
development of the spines is the same and in spio we find the same spines about the
middle of the ventral side of the costa. The situation in spio appears to be less dif-
ferentiated (in Fig. 36 indicated by 10a). It is most unlikely that only in geron geron
the original unspined condition was maintained, as this would contradict other characters
and the spined condition would have evolved independently in some or all other forms.
So we may assume that the condition in geron geron is apomorphous as compared with
the condition in the other species of the group (in Fig. 36 indicated by 10b).

In Fig. 36 the supposed evolution of the phlomidis group is summarized and the
plesiomorphous and apomorphous conditions of the characters dealt with are represented
symbolically.

The subspecific differentiation of phlomidis and doris does not need to be dealt with
here; see chapter 3 and, for doris, see also below.

2. Geographic history.

The species of the phlomidis group have a peculiar distribution: they occur in the
eremian parts of the Palaearctic. Although there exists some overlap, the phlomidis group
forms geographically a link between the sertorius group and the African Spialia species.
However, as will be shown below, the phylogenetic link between the sertorius group and
its African congeners has nothing to do with the phlomidis group.

In the last paragraph spio was shown to be the probable phylogenetic connection
between the phlomidis group and the African Spialia species. Moreover, spio appears to
have retained more characters of the supposed common ancestors than the species of the
phlomidis group. This does not necessarily mean that the phlomidis group has an
Ethiopian origin, it only makes probable that the development within the group is of a
rather recent time. From Fig. 36 it follows that the most recent development was the
separation of doris, osthelderi and geron, the most eremian species. This makes it prob-
able that the desiccation of the southern Palaearctic and the broadening of the eremian
zone in this region have played a part in the evolution of the phlomidis group, as the
common ancestor of doris, osthelderi and geron may have been forced to adaptations
to a very dry environment. However, once eremian, this supposed species must have
suffered more from wet than from dry periods. Thus, the Pleistocene Pluvial Periods (cf.
Moreau, 1955) have undoubtedly been important for the evolution of the phlomidis
group by causing geographic isolations. It is, however, impossible to date or to locate
more or less exactly the isolations that necessarily have started the evolution of the
phlomidis group. This is due to our very incomplete knowledge about both the phlomidis
group and the history of the regions concerned. Undoubtedly, the Zagros Mountains, the
Mesopotamian plains and the Nile Valley have at some time played a part in the

260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

geographic history of the phlomidis group, but this part is not clearly discernible, at least
at the specific level. The subspecific differentiation of doris and osthelderi, however,
appears to be clearly influenced and probably caused by a Pluvial Period that moistened
the Zagros Mountains and made it possible for forests to develop in the Mesopotamian
plains. Also an inundation of the latter region by transgression of the sea may have had
influence. Thus, we have a western and an eastern subspecies in osthelderi (ssp.
osthelderi and ssp. gecko, respectively) and a western group of subspecies and an eastern
subspecies in doris (ssp. doris + amenophis and ssp. evanida, respectively). It is con-
fusing that doris doris and doris evanida are so much alike and doris amenophis so
different, as one is easily inclined to suppose the closest relationship between forms that
resemble each other most closely. However, the large spots of doris amenophis are un-
doubtedly apomorphous; thus the resemblance between doris doris and doris evanida is
based on a plesiomorphous character. As clearly pointed out by Hennig (1966), cor-
respondence in plesiomorphous characters is no measure for phylogenetic relationship.

The distribution of doris amenophis suggests that the Nile is, and probably has been
in the past, a distributional barrier, as doris amenophis is not known from the area west
of the Nile. The occurrence of an isolated population of doris far to the west, in
Morocco (doris daphne) shows that doris once had a larger distribution in N. Africa.
As it is unclear why a species adapted to desert environment would die out in the desert,
the lack of specimens of doris from the region between Morocco and Egypt may be rather
the result of poor exploration than of true absence.

c. The sertorius group

1. Evolutionary history.

We will deal with the characters in the same way as in the foregoing paragraph.

(1). Valva: costal process. In Spzalia the costal process usually consists of a ventro-
distal thickening of the costa, with hairs or spines. This structure may be very dif-
ferentiated and is often roofed in by a fold from the dorso-distal part of the costa. The
type found in sertorius and its allies, viz., an appendage of the ventro-distal part of the
costa, set with spines that are upturned and directed to the dorso-distal part of the valva,
is not found elsewhere. Such a differentiated structure seems to be apomorphous rather
than plesiomorphous. The plesiomorphous stage of this character is difficult to indicate
in the other Spialia species, as most species appear to be rather differentiated. It is prob-
able that the plesiomorphous stage of the sertorizs type of costal process is no longer in
existence. The most important statement, however, is that the resemblance between the
species of the serforius group in this case is based on an apomorphous character. An
independent development of this character in various species in very unlikely.

(2). Valva. In the species of the sertorius group, and only in these species, the inside
of the valve is provided with a conspicuous, hairy fold. This must be an apomorphous
character as it is absent in all other Spialia species, as well as in related genera.

(3). Female genitalia: 8th abdominal segment. In the sertorius group the ventral and
dorsal sclerites of the 8th abdominal segment are fused together so that the sclerotization
of the segment consists of two parts that dorsally are connected by a membrane and
ventrally by a narrow sclerotized antevaginal band. In other Spialia species the 8th ab-
dominal segment bears four sclerites that are hardly or not fused together. The fusion
of sclerites can be taken as an apomorphous character.

R. DE JONG: Systematics and evolution of Spialia 261

(4). Gnathos. All species of the sertorius group have a well-developed gnathos. In
superspecies sertorius the strong differentiation into a stalk and a spined “head” appears
to indicate an apomorphous stage as compared with the slighter differentiation in mafa
and galba.

(5). Valva: costal process. If we imagine the evolution of the costa in Spialia from
a plain structure towards a structure with highly differentiated appendages, there can be
seen a gradual development of the costa in the sertorius group: from mafa through galba
to sertorius and orbifer the costal process is shifted as it were towards the distal end
of the costa and, eventually, replaces it. The difference between mafa and galba is slight
as compared with the difference between galba and sertorius. So we can call the sertorius
type the apomorphous stage and the mafa/galba type the plesiomorphous stage.

(6). Valva. The hairy fold at the inside of the valva shows different forms. There
appears to be a gradual change from mafa through galba and sertorius to orbifer as the
fold changes its position in that direction from horizontal-dorsal through vertical to
horizontal-ventral. Possibly the development of the fold has run parallel to that of the
costal process, but this cannot be more than an assumption as it is yet impossible to
indicate the apomorphous and plesiomorphous stages of the position of the fold. Thus,
we cannot use this character for the reconstruction of the evolution of the sertorius
group.

(7). Tegumen and uncus. In mafa and galba there is an area between the uncus and
the tegumen that is only weakly sclerotized, thus forming a more or less triangular
“fenestra”. In superspecies sertorius this ‘‘fenestra” is absent, though the connection
between the uncus and the tegumen is clearly visible. The phylogenetic importance of this
“fenestra” is still obscure. It may indicate that mafa and galba are closely related, as it is
possible that the “fenestra” is an apomorphous character, but this statement is not very
well founded at the moment. If also other characters are included, we may suppose that
it is desirable to unite mafa and galba into a superspecies. This, however, falls outside
the scope of this paper.

(8). Colour of the underside of the hind wing. In Spzalia this underside shows
various colours, mostly greenish or ochreous. In sertorius the colour is red or reddish
ochreous, in orbifer it is greenish, but in orbifer carnea it is as red as in sertorius sertorius.
The red and reddish ochreous tinges seem apomorphous if compared with the greenish
colour. However, the red colour must have originated independently in sertorius and in
orbifer carnea, as other characters (e.g. position of the hairy fold of the valva, measure-
ments, spotting) show that orbifer carnea is an orbifer lugens with a red underside of
the hind wing.

We can also suppose a parallel development of the red colour in the subspecies of
sertorius, but this assumption is not supported by other characters (e.g. hairy fold,
spotting). So we may assume that the red colour originated twice. In Fig. 37 this is
indicated by the numbers 8a and 8b.

(9). Spotting on the underside of the hind wing. This is very variable in Spzalia,
some species having distinct spots, others straight or curved bands. If there is a distinct
central band, this band is directed to or conjoined with the outer spot in space 7 or it is
directed to a point between the inner and outer spots in space 7. It is, however, directed
to or connected with the inner spot in space 7 in sertorius and orbifer and in phlomidis.
As stated above, the situation in phlomidis can be considered apomorphous. Also the
situation in superspecies sertorius can be called apomorphous. Apparently the resemblance

262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

between phlomidis and superspecies sertorius in this respect is based on a parallel
evolution. In Fig. 37 the character of the direction of the central band is indicated by
the number 9a.

A main differentiating character between sertorius and orbifer is the shape of the
central spot on the underside of the hind wing. This spot is angular in sertorius and
rounded in orbifer. If a continuous central band is broken into spots by darkening of the
veins one expects that the central spot is angular along veins 4 and 6, while these angles
may become rounded off. This appears to have been the case in superspecies sertorius,
but in sertorius the central spot became still narrower along veins 4 and 6 and, at the
same time becoming angular along vein 5. I cannot consider this character otherwise than
apomorphous. In Fig. 37 it is represented by the number 9b.

=) Gh Tl Q un > a
oe 3 Ss EC a DRE:
>. = D 2 Sus
wie! ij Den, a aves
a 8 eS TR
a Oo u

HH \ =)

12 b

Oma OO
on
O

@ (Ze

OOS
UI

Gore
Oene
N

Fig. 37. Supposed evolution in the Spialia sertorius species group. The numbers indicate characters
as indicated in the text. O = plesiomorphous condition, ® = apomorphous condition

R. DE JONG : Systematics and evolution of Spialia 263

The most advanced development of the central spot is found in sertorius ali, where
the spot is projected towards termen and base, while also the submarginal spots are
stretched out. This situation has been represented in Fig. 37 by the number 9c.

(10). Colour of the spots on the upperside. In all Spzalia species this colour is white,
but in sertorius therapne and in the summer brood of sertorius ali it is yellow. Un-
doubtedly the yellow colour is an apomorphous character and there is no apparent need
for considering the yellow colour to have evolved independently in sertorius therapne
and sertorius ali. It would, however, be very elucidating if the genetic background of this
character was known, as apparently ecological factors influence the expression of the
character (at least in sertorius ali).

(11). Submarginal markings on the upperside. The development of the submarginal
markings is subject to much variation in superspecies sertorius. Though the genetic back-
ground is rather obscure, the expression of this character appears to be determined more
ecologically than geographically, see under sertorius sertorius and orbifer orbifer. How-
ever, the whole Asiatic range of orbifer east of the Caspian Sea is inhabited by pop-
ulations in which the submarginal spots are faint or absent, thus indicating that these
populations are apparently incapable of developing well-marked submarginal spots. As
in almost all Spzalia species the submarginal spots are well-developed, the absence of
these spots in some populations of a single species appears to be an apomorphous
character.

(12). Number of broods per year. In superspecies sertorius the usual number of
broods per year is two, though one brood may be more or less suppressed by ecological
factors (mainly at higher elevations in the mountains). However, in spite of the large
ecological diversities in Asia, the populations of orbifer east of the Caspian Sea appear
to be single-brooded. Unless we assume that the double-brooded condition arose in-
dependently in sertorius and in the orbifer populations west of the Caspian Sea, the
single-brooded condition must be considered apomorphous. In Fig. 37 this situation is
represented by number 12a.

Very probably sertorius therapne also is single-brooded. In this case, too, it must be an
apomorphous condition, that evidently arose independently of the development in Asia
(Fis-37:12b).

2. Geographic history.

It is striking that there is only a single Spialia species in the Oriental region (galba)
and, apart from the eremian phlomidis group only two Sptalta species (united in super-
species sertorius) in the Palaearctic region, while there are many species in the Ethiopian
region. Together with the supposed evolutionary history as expressed in Fig. 37, this
makes it probable that galba and superspecies sertorius have an Ethiopian origin.
Evidently, superspecies sertorius is the older offshoot. As the ancestor of superspecies
sertorius appears to have been the only one that successfully colonized the Palaearctic,
the eremian zone between the Palaearctic and Ethiopian regions must have been a serious
barrier since long, even when it was much narrower than at present. In this connection,
it is interesting that the eremian zone not only did not function as a barrier for the
ancestor of the phlomidis group, but even became its habitat.

Possibly the widening of the eremian zone quickened and intensified the geographic
isolation of the ancestor of superspecies sertorius. The evolutionary history and the little

264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

variation in Asia east of the Caspian Sea show that this part of the range was colonized
rather recently. Thus, the ancestor of superspecies sertorius inhabited the Mediterranean
and there it became differentiated in sertorius and orbifer. The distribution of these
(semi-)species shows the apparent cause of the differentiation, viz., geographic isolation
in a western (sertorius) and an eastern (orbifer) Mediterranean refugium during a
Glacial Period. The occurrence of orbifer in Sicily indicates that sertorius most probably
originated in Spain, in a later time invading Italy where orbifer had become extinct
during the preceding Glacial Period (possibly except in southern Calabria).

The close relationship between serforius therapne and sertorius ali is somewhat
puzzling as their distribution areas lie far apart, while the intermediate area is populated
by sertorius sertorius. If we have interpreted the facts correctly, the following explanation
may be an exact reconstruction of the history. At a moment that the red colour of the
underside of the hind wing had developed to a certain degree, but probably not yet to
the bright red of some recent populations of sertorius sertorius, in the western-most
populations the white spots of the upperside became yellowish in the summer brood,
while in more eastern populations (Italy) the spots remained white. Possibly this dif-
ferentiation was initiated and influenced by a geographic discontinuity caused by a Glacial
Period, but this is, of course, not necessary. A Glacial Period destroyed the greater part
of the western-most populations and west of Italy the species survived in isolated areas
only, viz., N. Africa, Corsica and Sardinia. As they are surrounded by water, these areas
could not have much importance for the expansion of the species in an Interglacial Period
and so the intermediate region became populated from another refugium (Italy). By
inbreeding and other factors, the populations of Corsica and Sardinia became single-
brooded without losing the yellow colour of the spots on the upperside: sertorius therapne
is, as it were, the summer brood of sertorius ali, without producing a spring brood.
Isolated in N. Africa sertorius ali, or rather its precursor, developed the peculiar markings
on the underside of the hind wing, but the yellow spots remained the characteristic of the
second brood.

There are two other explanations possible for the discontinuous distribution of the
yellow spots: (1) the Iberian populations were not destroyed, but they lost again the
newly developed character of the yellow spots, and (2) the yellow spots arose in-
dependently in sertorius therapne and sertorius alt. Both explanations appear to me less
probable than the one given first.

The great diversity of sertorius in Spain may be an argument for the assumption that
Spain was colonized from the east by a multiple invasion so that its genetic variability did
not yet attain an equilibrious condition. In this way, possible remnants of populations
with yellow spots in the second brood could have been trodden under foot.

In the meantime, orbifer invaded Asia. The lack of data from E. Russia and W. Siberia
makes it impossible to decide about the route: north or south along the Caspian Sea. The
small degree of variation over the large Asiatic range is surprising. It can be the result
of a rapid expansion. However, the differentiation between orbifer lugens and orbifer
carnea seems the result of geographic isolations in glacial refugia, so the presence of
orbifer east of the Caspian Sea must date from the last Interglacial or earlier. Therefore,
if the small amount of variation is the result of a rapid expansion, it must have been
orbifer lugens that expanded its range rapidly from a glacial refugium. In this connection
it can have been advantageous, that orbifer lugens is single-brooded, giving it the highest
survival rate in the extreme continental climate.

R. DE JONG : Systematics and evolution of Sptalia 265

The location of the refugium of orbifer carnea is not difficult, as this subspecies
evidently did not extend its refugial range very much. So it must have survived the last
Glacial Period in the Afghan refugium (cf. Gross, 1961; De Lattin, 1967). For orbifer
lugens it is much more difficult to indicate a glacial refugium. From its present distrib-
ution we may conclude that it did not occur south of the Hwang-Ho and that the
Mongolian-Siberian refugia are less probable for orbifer lugens (species extending from
these refugia usually have another distribution pattern). So the Turkestanian and Man-
churian refugia are considered for glacial refugia of orbifer lugens. The former seems
the most probable, as otherwise orbifer lugens would already have colonized the whole of
its present range before the Last Glacial. In that case, we would have expected a further
differentiation of the species in Asia.

The differentiation between orbifer orbifer and orbifer hilaris is undoubtedly young,
not older than since the Last Glacial. Presumably, o. orbifer originated in a refugium in
the Balkans and W. Turkey, while orbifer hilaris did so in Syria/Iran. See, however, the
note under ssp. hilaris in Chapter 4.

Summarizing, the geographic history of superspecies sertorius may be represented as
follows.

(1). A northern branch of an Ethiopian species became isolated in the Mediter-
ranean.

(2). Forced by advancing deterioration of the climate during a Glacial Period, the
Mediterranean population became divided into a western and a more eastern group
giving rise to sertorius and orbifer, respectively.

(3). During a following Glacial Period, orbifer died out in Italy, but could survive
in Sicily. In the next Interglacial orbifer did not succeed in regaining the lost Italian
territory, but was replaced there by sertorius. It is interesting to note, that the isolation of
orbifer in Sicily did not lead to the development of a recognizable differentiation.

(4). In the western populations of sertorius the spots became yellow in the summer
brood. A following Glacial Period destroyed all or almost all sertorius populations in
Spain. The character of the yellow spots could survive in Corsica-Sardinia and NW.
Africa only.

(5). The “empty” Iberian peninsula was invaded at least once and probably twice
or more times by sertorius from Italy, resulting in much variation, that is partly locally
determined.

(6). In the meantime, orbifer invaded Asia east of the Caspian Sea, where it became
single-brooded during isolation in a glacial refugium, possibly in Turkestan. After
extending its range during an Interglacial, the Central Asiatic population was driven back
to glacial refugia during a following Glacial, in Afghanistan (orbifer carnea) and
possibly in Turkestan (orbifer lugens).

(7). In the eastern Mediterranean a differentiation took place between orbifer orbifer
and orbifer hilaris in glacial refugia in the Balkans and W. Turkey and in Syria/Iran,
respectively.

(8). After the Last Glacial sertorius sertorins extended its range from Italy and
presumably also from Spain to the north and the east. In Central Europe it encountered
orbifer, which advanced from its Balkan refugium to the north and west. The greatest
extension of the range was made in Asia by orbifer lugens that from its presumably
Turkestanian refugium colonized Asia as far as the Amur region and NE. China.

The geographic history outlined above clearly shows the impact of the Glacial Periods

266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

on the development of the superspecies sertorius. It is, however, difficult to date the
various geographic isolations that must have taken place, as the influence of the different
fases of the Glacial Periods are still little understood. Possibly various fases lasted too
briefly to have much influence. But even if we claim a whole Glacial Period for each
of the supposed geographic isolations, it is still obvious, that the entire development
of superspecies sertorius took place during the Pleistocene. In other words, the super-
species Spialia sertorius originated as a result of the climatic fluctuations of the
Pleistocene.

A few words must be said on an earlier hypothesis on a part of the geographic history
of the superspecies sertorius. To explain the occurrence of species in islands authors have
often projected land connections between islands and continents. Even if these connections
are geologically demonstrable, we cannot be sure of the use the species made of it. An
author who often used former land connections for explaining butterfly distribution in
Italy, is Verity (1940). He considered the occurrence of orbifer in Sicily the result of the
presence of a land connection from Greece through Calabria and Sicily to Tunesia in
the Miocene. Apart from the question whether the connection existed in that form,
there are the following objections against this explanation:

(1). It does not explain the presence of sertorius instead of orbifer in Calabria.

(2). It does not explain, why it is not sertorius ali that occurs in Sicily.

(3). The Miocene is much too long ago to explain the recent differentiation at the
subspecific or semispecific level.

According to Verity, sertorius ali is “la forma più primitiva”, from which sertorius
sertorius, sertorius therapne and orbifer originated directly, but in different periods. In
this hypothesis orbifer is a “trasformazione antichissima”, sertorius sertorius originated
“fin dal Miocene” and sertorius therapne is a “derivazione recente”. Unfortunately, the
only argument given by Verity is the shape of the spots on the underside of the hind
wing; this argument is far from being convincing. If orbifer invaded Sicily in the
Miocene, its differentiation from sertorius ali must have taken place earlier. It is quite
incredible, that a differentiation originating so far back in the Tertiary could maintain
itself rather undisturbed in a region that underwent enormous ecological changes.
Reasoning in the way of Verity and various other authors, who neglect the overwhelming
effect of the Pleistocene Glacial Periods on the differentiation and distribution of the
species and forget that the Miocene ended some 12 million years ago (i.e. 12 times as
long ago as the beginning of the Pleistocene), would lead to the assumption that the
Hesperiidae originated in the beginning of the Tertiary or even earlier and that the
Lepidoptera originated tens of millions of years before there were flowers, to suck honey
from.

Summarizing, the explanation given by Verity does not explain anything. Apparently
the presumption that butterflies cannot cross stretches of sea, is incorrect. This is also
demonstrated by the discovery of sertorius sertorius in Corsica. This subspecies apparently
reached Corsica in recent times, without the help of a land bridge. Moreover, Moreau
(1952), summarizing the palaeogeography of Africa, did not mention a Greco-Tunisian
land bridge in the Miocene, but only a Sicilo-Tunisian one in the Pliocene. This is much
more in accordance with our hypothesis, as it places the evolution of superspecies sertorius
after the severing of the Sicilo-Tunisian land bridge, i.e. entirely in the Pleistocene.
Otherwise, sertorius ali would occur in Sicily, or orbifer in NW Africa.

Palaearctic species ultimately originating from Ethiopian species appear to be rare.

R. DE JONG: Systematics and evolution of Spialia 267

Their scarcety illustrates the effectiveness of the desert belt (partly in combination with
the Mediterranean) as a barrier, though the width of this belt has varied much. Among
the Palaearctic Hesperiidae the two Gegenes species possibly have an Ethiopian ancestry,
but they are much more confined to a dry, steppe-like habitat than Spialia sertorius and
orbifer.

Undoubtedly other examples can be found among the Palaearctic Lepidoptera, but I do
not know of such cases from the literature. Professor Dr. K. H. Voous kindly informed
me that a comparable case exists among birds, viz., the Collared Turtle Dove, Streptopelia
decaocto, of the roseogrisea/capensis group. But at least part of the extension of this
species into the Palaearctic is very young, not older than some centuries.

6. SUMMARY.

1. Four species and one superspecies, consisting of two (semi)species, of Spialia,
have been recognized in the Palaearctic Region. They can be assigned to two species
groups, viz. the phlomidis group and the sertorius group. The phylogenetic relationship
between these groups has not been discussed in this paper, for an examination of such a
relationship a revision of the Ethiopian Spialia species is necessary.

2. The phlomidis group (four species) is confined to the eremian parts of the
southern and central Palaearctic. Phylogenetically it is directly related to the Ethiopian
species Spialia spio.

3. The serforius group has a very wide distribution. In the Palaearctic, where it
comprises a superspecies with two (semi)species (sertorizs and orbifer), it occurs from
the Atlantic to the Amur region and China. An Ethiopian (mafa) and an Oriental species
(galba) must also be assigned to this species group.

4. By the application of a subspecies definition that uses geographic isolation (at
present or formerly) as criterion, several subspecies distinguished by previous authors
have been assigned to clinal (local) and nongenetic ecophenotypic variation.

5. The theoretical basis for the arrangement of the species and subspecies adopted in
this paper is the discussion on the evolutionary and geographic history. Several characters
have been examined as for their apomorphous or plesiomorphous conditions.

6. The phlomidis group possibly has an Ethiopian ancestry. It developed in a recent
time, probably not before the Pleistocene, in the eremian parts of the Palaearctic. The
history of the group is obscured by lack of data.

7. The superspecies sertorius is the Palaearctic off-shoot of an Ethiopian ancestor
which also gave rise to an Oriental representative of the serforius group. Superspecies
sertorius originated in the Mediterranean as a result of the Pleistocene climatic changes.
East Asia was probably reached only in postglacial times, from Turkestan.

268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

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270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 6, 1974

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Postscript

After having sent this paper to the printers’, I came across a recent article by L. G.
Higgins (J. Ent. (B) 43 (1) : 83—87) on a new subspecies of Spzalia phlomidis from
the Lebanon, which he named S. phlomidis kiki. Evidently, this name is a junior synonym

of Spialia phlomidis hermona Evans.

abscondita 257
adenensis 241
ali 247
alioides 251
amenophis 240

carnea 256

daphne 240
doris 239

evanida 241
eucrate 251
eupator 234

galba 230, 246, 261
gavarniensis 251
gecko 236

Gegenes 266

geron 238

gracilis 250

Index
guadarramensis 251

hermona 234
hibiscae 246
hilarıs 254

lugens 255

mafa 230, 246, 261
melotis 236

minor 250, 253
murasaki 253
Muschampia 228

nanus 257
nomas 236

orbifer 252
osthelderi 235

parvula 250

phlomidis 233
Platygnathia 228
posttesselloides 254
Pyrgus 228, 236

sao 228, 246
secunda 255
sertorius 244, 246
Spialia 228

spio 256, 257, 259
struvei 239
struveoides 236
subgracilis 250
Syrichtus 228, 236

tesselloides 254
tessellum 236
therapne 248
therapnoides 247

zebra 228

oa ee

I»

No. 4. F. Willemse, 1968. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov
and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl.
D.FI. 25.—. È i i

No. 5. C. A. W. Jeekel, 1970. — Nomenclator generum et familiarum Diplopodorum, 412 pp.
D.FL.100.—.

No. 6. J. G. Betrem, 1971. — The African Campsomerinae (Hymenoptera, Scoliidae), 326 pp.,
47 figs., 6 pls. D.Fl. 120.—.
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THE GRAVENHORST, SCHISDTE AND FOERSTER TYPES
BELONGING TO THE GENUS MEGASTYLUS SCHIÖDTE, 1838,
WITH KEYS TO THE SPECIES (HYMENOPTERA,
ICHNEUMONIDAE, MICROLEPTINAE)

by
G. VAN ROSSEM

Plant Protection Service, Wageningen

ABSTRACT

The lectotypes of Plectiscus flavopictus Gravenhorst, 1829, Megastylus cruentator Schigdte, 1838,
Megastylus mediator Schiode, 1838, M. pumilio Foerster, 1871, Myriarthrus cingulator Foerster,
1871, and Dicolus pectoralis Foerster, 1871, are designated. The male of Megastylus cruentator
Schigdte is described for the first time. Two subgenera of Megastylus are recognized: Megastylus
and Dicolus Foerster, 1868. Some new combinations and synonymy are proposed. The paper is
illustrated with figures and 4 photographs.

TAXONOMY AND NOMENCLATURE

Foerster’s (1868) establishment of Myriarthrus was incorrect as for no obvious reason
he rejected the name Helictes Haliday, 1837. As the type species of Myriarthrus, Foer-
ster (1871) took Ichneumon erythrostoma Gmelin, sensu Gravenhorst, 1829, the males
of which have concavities on certain flagellar segments. Thus Foerster misunderstood
Gravenhorst’s description and consequently Myriarthrus became an unnatural assembly
of species. Viereck therefore proposed Myriarthridea with Myriarthrus cingulator Foerster
as the type species. The generic names Dicolus Foerster, 1868, and Myriarthridea Viereck,
1914, were both placed in the synonymy of Megastylus by Townes (1971).

There is sufficient reason to recognize Dicolus Foerster as a useful taxon, namely as a
subgenus of Megastylus. The two subgenera of Megastylus are separated in the keys
following below. The deep propodeal fossa and the very slender postanellus seem to be
well defined characters of Dicolus.

I have not searched the Foerster collection for concealed Megastylus material in the
modern sense. The Foerster (1868 and 1871) “Plectiscoid’’ genera were actually first
revised by Perkins (1962) and later rearranged by Townes (1971).

SEPARATION OF THE SEXES

It is difficult to determine the sex of many specimens at first glance. For this reason
Schigdte and later authors never recognized the proper male of M. cruentator. What
Schigdte thought to be males of M. cruentator turned out to be females. I found this by
making microscopic preparations of genitalia of the syntypes (PI. 1 Fig. 2). As a con-
sequence the male of M. cruentator was never described. One purpose of this paper is to
rectify this strange confusion.

273

274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 7, 1974

The sexes in Megastylus are easily separated by the shape of the antennal bristles.
Males have very short, close and vertical setae (Fig. 1); the female antenna is characterized
by the much longer and scarcer pilosity (Fig. 2).

Fig. 1—4. Antennae and legs of Megastylus. 1, M. cruentator, 4, Planken-Wambuis near Ede,
15.V.1966 (coll. van R); 2, the same, ©, N Spessart, Lochmühle, 17-22.X.1971, trap (coll. van R);
3, M. pectoralis, ®, left hind tibia (inner side), Lousberg (Aachen), 15.X., coll. Foerster (ZSM);
4, M. impressor, 2 , holotype, right hind tibia (inner side) (MC).

À NOTE ON THE FOERSTER TYPES

The Foerster Megastylus, Myriarthrus and Dicolus material is quite undisturbed, but
it has a remarkable feature, namely, some kind of discoloration. Most specimens have a
light brownish tone of the integument which is not the natural colour. The types of
many species are surprisingly small specimens. Mounting and labelling are excellent.

TENTATIVE KEY TO FEMALES

1. Postanellus short, index 3.5—5.0. Subgenus Megastylus. . . . . . .. 2
— Postanellus long and slender, index 6.0—10.0. Subgenus Dicolus . . . . 5
2. Second gastral tergite striate. Hind coaxe on the lower inner side granulated*). In

typical specimens frontal orbits always with broad white lining. Mesoscutum with

*) Difficult to see.

2»

G. VAN ROSSEM : The types of Megastylus 275

anterolateral yellowish marks, from these yellowish lines converge towards and coalesce
on disk. Episternum 2 always completely black. Third gastral tergite in most cases
yellowish to brownish . . . . Jet flavopictus

Second gastral tergite with fine nde microsculpture, oi never striate 3

Frontal orbit (above antennal sockets) with conspicuous white lining. Rather small
specimens, 3.5—4.6 mm, difficult to e from M. piane ve rufipleuris

Frontal orbit without white lining. . . RAA HART A

Middle femur stout, index 3.6—3.8. Small to te neels DI el, 2 mm. Face,
frons and thorax without any white or yellowish marks. Antennal base and legs,
including coxae light brownish. Head, thorax and gaster almost uniformly dark
reddish brown*) to black, except for yellow marking of gastral segment 3. All
femora stout, indices: 3.0—3.1 (1); 3.6—3.8 (2); 5.0 (3). . . . . pumilio

Middle femur slender, index 5.1—7.0 Specimens with strong variability in size
(4.0—7.0 mm) and colour, the latter ranging from deep black: head (exept
clypeus), thorax (except legs) and gaster to specimens with most of notum 1,
episternum 2, mesosternum and pleural parts of propodeum fulvous to deep brick-
red. Palpi, mandibles and clypeus often white to dirty white. Anterior margin of
notum 1 often marked white. ent with lighter gastral tergites 2 and 3 occur

. cruentator
Uni a TI real in ihe 0. 7 proximal ape aus causing a con-
striction in the 0.3 distal part (Fig. 3). . . . ME
Hind tibia weakly inflated in proximal part, Das not anal ci in the
distal part (Fig. 4) . . . . e 7/
Rather small specimens (3. gal 4 a) Eion chi. dn acri of antennal
insertions without a broad yellow spot. . . . . . . subtiliventris

More robust specimens (5.2—5.5 mm). Frontal orbits alse: laterad of antennal
insertions with broad yellow spot. Mesoscutum with vague yellow lines, coalescing
Omdisk … . 5 . . pectoralis

Basitarsus of front Teg avon 0. 56 gi Losi ont a Eu sans (and also
following tarsal segments) of front leg strikingly slender. Antennal cleaner large,
causing conspicuous outcurving of basitarsal base (Fig. 5). Hind femur not inflated
in proximal part. . . u... .excubitor

Basitarsus of front leg ben 0. 330. 43 Of Cosi front sn not slender. Anten-
nal cleaner not conspicuously outcurved (Fig. 6). Hind femur somewhat inflated in
Prosa part NEN N a) OR, A NE SENS im pressof

TENTATIVE KEY TO MALES

Males of M. rufipleuris, M. subtiliventris and M. excubitor are unknown

Hind tibia somewhat inflated in its proximal 0.67 part (Fig. 7) (subgenus Dcolus)

3 M. pectoralis and perhaps M. subtiliventris

iad tibia not fred in ies i Parto) AE hace ED
First and second gastral tergite striate. Specimens abundantly i with yellow:
palpi, mandibles, malar space, clypeus, face entirely, frontal orbits, pleuron 1, prono-
tum, mesoscutum with broad anterior lining and parallel lines coalescing on disk.

*) Especially in museum specimens.

276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 7, 1974

Episternum 2 for the greater part black. Genitalia, Pl. 1 Li 3. . M. flavopictus
— First and second gastral tergite not striate. . . . Me,
3. The following characters combined: propodeum us a Hs transverse

impression (fossa) in its 0.25—0.28 proximal part, thus clearly away from the
apical boundary of notum 3 (postscutellum) ; postanellus relatively long and slender,
index 8.8—12.5*); hind femur long and slender, index din A Hind tibia slimly
clavate (Figs8)) MCO. >... M.impressor
— Propodeum with either a ee Ga M. oe) or none evident impression; posta-
nellus less slender, index < 8.8 (5.5—8.0); hind femur of normal Du: index
5.8—7.0. Hind tibia weakly to evidently clavate. . . 4
4. Minute specimens (about 2.7—3.2 mm long). Index alle DE sig 5. Thorax
entirely brown or black. Hind tibia somewhat clavate towards apex. Legs including
coxae yellow. Face brown (to yellow?). . . . . … … M. pumilio
— More robust specimens, 4.1—5.4 mm long, adat m with yellow: palpi,
mandibles, clypeus, malar space, face entirely, frontal orbits, pleuron 1, pronotum,
episternum 2. Vertex and back of head black. Mesoscutum for the greater part black,
with vague brownish lining on disk. Propodeum including sternal parts mostly black.
Index postanellus 6.0—8.0. Hind tibia clearly clavate towards apex. Legs with all
coxae: yellow. Genitalia; Pl. 1. Fig. 4 CN 22... Mi cruentatof

Subgenus Megastylus Schigdte, 1838

Type species: M. cruentator Schigdte, 1838.
Subgenus Megastylus: Thomson, 1888, Opusc. Ent. 12 : 1311.

Megastylus cruentator Schigdte

4 (! 2) Megastylus cruentator Schigdte, 1838, Rev. Zool. Cuv. 1 : 139.
Megastylus cruentator: Schigdte, 1839, Mag. Zool. (2)1 : 4-5.
Q Megastylus mediator Schiodte, 1838, Rev. Zool. Cuv. 1 : 139.
Megastylus mediator: Schigdte, 1838, Mag. Zool. (2)1 : 5
® Megastylus nigriventris Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 105. New synonymy.
9 Megastylus conformis Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 105. New synonymy.
Q Megastylus fuscicornis Foerster, 1871, Verh. naturh. Rheinl. 28 : 105. New synonymy.
Megastylus cruentator: Townes, 1971, Mem. Amer. Ent. Inst. 17 : 205.
Megastylus mediator (= cruentator): Townes, 1971, Mem. Amer. Ent. Inst. 17 : 205.

Characteristics of the type specimens. — M. cruentator. Female. Front wing 4.9**)
Labels: a white printed museum label “Danmark ex coll. Schigdte’’; lectotype label and
Q added by me; distal part of gaster cut off and micro-slide for genitalia fixed on pin.
M. cruentator det. van R.

M. mediator. Female. Front wing 4.4 mm. Labels: a white label (probably original)
“© Kjöbenh. [the type locality} Schiodte”; a white museum label “Danmark ex coll.
Schigdte”’; lectotype label by me: M. cruentator det. van R.

M. nigriventris Foerster. Female. Front wing 5.7 mm. Labels: a white label (original)
“45. gl.***) Mont-joie”; a blue-rimmed box label “nigriventris Frst.”. Holotype labelled
by me: M. cruentator det. van R.

*) Based on 3 specimens and probably not quite reliable.
**) From base of costal margin.
***) Meaning: “Glieder’’ = segments of antenna.

G. VAN ROSSEM : The types of Megastylus 277

M. conformis Foerster. Female. Front wing 5.0 mm. Labels: a white label (original)
‘45 gl Aachen"; a blue rimmed box label “conformis Frst’’. Holotype labelled by me:
M. cruentator det. van R.

M. fuscicornis Foerster. Female. Front wing 3.2 mm. Labels: a white label (original)
“9 + 39 gl. Aachen”; a blue rimmed box label “fuscicornis Frst.’’. Holotype labelled
by me: M. cruentator det. van R.

Description of the male. — Body length 4.0—5.4 mm. Front wing 3.0—4.0 mm.
Head with the following parts yellow: palpi, mandibles, lower gena, clypeus, entire face,
lower part of frons and part of inner orbit. A sharply bounded rectangle below the
central ocellus browish to black. Vertex and back of head black. Eyes and ocelli large.
Labial palpi short; maxillary palpi strikingly long and slender, reaching at least well
beyond front coxa. Mandible short, somewhat swollen at base, strongly tapering towards
apex; teeth lying in horizontal plane. Clypeus strongly convex with characteristic, erect,
sparse hairs. Clypeal fovea deep, with an open anterior tentorial pit. Malar space wide,
with a groove. Head preponderantly polished, here and there finely coriaceous. Anten-
nae, slender, of body length.

Thorax characteristically coloured: entire pronotum, pleuron 1, all coxae, episternum 2
(mesopleurae) and mesosternum ivory yellow towards reddish brown. Mesoscutum
entirely black, in some specimens with light brown streaks along the prescutal sutures
which may coalesce on the disk and backwards towards the scutellar fovea. Scutellum
often brownish. The entire propodeum including the sternal part mostly black. Sculpture
of pronotal and mesopleural (episternum 2) parts almost polished. Mesoscutum strongly
convex, very finely coriaceous, prescutal sutures (notauli) weak. Propodeum only with
pleural carina, sometimes posterior transverse carina present; dorsally finely coriaceous
with a vague tendency towards transverse striation. Prepectoral carina present, in most
specimens not reaching suture between notum 1 and episternum 2. Entire thorax with
widely placed suberect, light-coloured hairs, wrich become more dense on the sublateral
and pleural parts of propodeum. Wings hyaline. Legs long and slender, yellowish, except
fuscous apex of hind tibia. All femora slender; index hind femur 5.8—7.1.

Gaster with 1st and 2nd tergite alutaceous and fuscous; 3rd and 4th tergite very
often yellow and finely alutaceous. Following segments mostly fuscous. Male genitalia as
in PI. 1 Fig. 4.

Female. — I refrain from giving a full description, as the female is sufficiently
characterized by the details given in the key. Two differential characters are: the slender
middle femur, index 5.1—7.0 and the absence of a white lining on the inner orbits of
the eyes. Ovipositor, Pl. 1 Fig. 2.

Remark. — In spite of the fact that Schiodte (1838, 1839) mentioned that he had males,
the type series of this species comprises females only. Nor is it clear from his description
that Schigdte recognized the male. The same holds for his second species (M. mediator)
as well as for Foerster’s three species, M. nigriventris, conformis and fuscicornis, and for
all females and all specimens of M. cruentator. This strange state of affairs was thus
pursued by Foerster (1871) and later by Schmiedeknecht (1908-1911). I conclude that
the male of this rather common species has never been properly described. When study-
ing the related M. flavopictus, I noted the strong sexual dimorphism in this species and
assumed that also in M. crzentator the male could hereby be recognised. A series of males
in the available material, formerly perhaps taken for M. flavopictus males, represent the
missing male of M. cruentator. It is described above.

278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 7, 1974

Biology. — The collecting dates on the examined material give some evidence that
two generations per year occur. The first from April to early June, the second from
August to the middle of October.

Material examined. — Denmark: 10 9, lectotype and 9 syntypes of M. cruentator,
Sielland (coll. Schigdte) (MC); 9, lectotype of M. mediator, Kjöbenhavn and 7 other
® , no locality (coll. Schigdte) (MC); &, as M. lineator!, Mon (coll. Schigdte) (MC).
Germany: @, holotype of M. conformis Foerster, Aachen (coll. Foerster) (ZSM); 9,
holotype of M. nigriventris Foerster, Montjoie (?) (coll. Foerster) (ZSM); ©, holotype
of M. fuscicornis Foerster, Aachen (coll. Foerster) (ZSM); 9, ND Spessart, Loch-
mühle, 17—22.X.*)1971, trap (coll. van Rossem). Italy: 9, Bolzano, 1914, leg. Smits
van Burgst (ELW). Netherlands: g', Ede, Sijsselt, 25.VIII.1963, trap (coll. van Ros-
sem); dg, Ede, Sijsselt, 7.IX.1963, trap (coll. van Rossem); 3 g', Ede, Planken Wam-
buis, 15.V.1966 (coll. van Rossem); 9, Ede (on window), 6.VII.1967 (coll. van Ros-
sem); 2 9, Ede, Sijsselt, 14.VIII and 28.IX.1970, trap (CJZ); 4 ®, Ede, Sijsselt, 12
& 22.V., 1 & 9.VI.1971, trap (CJZ); 9, Radio Kootwijk (Gerritsfles), 27.IX.1964
(coll. van Rossem); g', Radio Kootwijk (Assel), 12.1X.1963 (coll. van Rossem); d',
Kemperberg**) 28.V.1967, trap (coll. van Rossem); 9, Hilversum, 9.IX.1960, leg. den
Hoed (KWR); &, Nuth (Limburg), 17.V.1968, leg. den Hoed (KWR); 9, Vaals
(Limburg), IV.1914, leg. Smits van Burgst (ELW). No locality; 2 9, leg. Schmiede-
knecht (coll. Smits van Burgst) (ELW).

Megastylus flavopictus (Gravenhorst) new combination

& Plectiscus flavopictus Gravenhorst, 1829, Ichn. Eur. 2(2) : 983.
4 (1) Megastylus lineator Schiodte, 1838, Rev. Zool. Cuv. 1 : 139. New synonymy.
Megastylus lineator: Schigdte, 1839, Mag. Zool. (2)1 : 7.
Megastylus lineator: Foerster, 1871. Verh. naturh. Ver. Rheinl. 28 : 104.
& ® Myriarthrus cingulator Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 103. New synonymy.
Myriarthrus flavopictus: Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 103.
Myriarthridea cingulator: Townes, 1971, Mem. Amer. Ent. Inst. 17 : 205.

Characteristics of the type specimens. — Plectiscus flavopictus. Male. Front wing 3.6
mm, 2nd tergite striate. Labels: none. Lectotype label added by me: M. flavopictus det.
van R. Probably the specimen from Genova (“sexus incerti").

Megastylus lineator. Male. Front wing 3.8 mm. Labels: a white, rimmed label (prob-
ably original) “Q Strandm (Strandmöllen) Drewsen”; a printed museum label “Dan-
mark ex coll. Schiedte”. Holotype label and & added by me: M. flavopictus det. van R.
Schigdte indicated the sex of this specimen erroneously.

Myriarthrus cingulator. Male. Front wing 3.0. Labels: a white label “G.32 gl.
Aachen”; a blue rimmed box label “cingulator Frst.”. Lectotype label added by me; 12
syntypes labelled paralectitype, M. flavopictus det. van R.

I refrain from giving full descriptions of both sexes because the differential characters
of this species are recapitulated in the keys. Genitalia g' (PI. 1 Fig. 3).

Material examined. — Austria: 9, Carinthia, Ostkarawanken, Ebriach, 580—750 m,
21—29.VII.1964 (coll. van Rossem). Denmark: g', the holotype of M. lineator, Strand-

*) All specimens from this locality were wrongly labelled: IX.
**) Near Arnhem.

G. VAN ROSSEM : The types of Megastylus 279

Fig. 5—8. Legs of Megastylus. 5, M. excubitor, right front leg, female, holotype (ZSM); 6, M.

impressor, right front leg, female, Strandmollen, coll. Schigdte (MC); 7, M. pectoralis, left hind tibia

(outer side), male, Aachen, coll. Foerster (ZSM); 8, M. impressor, left hind tibia (outer side),
male, Viborg, coll. Schigdte

möllen, leg. Drewsen (coll. Schigdte) (MC). Germany: &, lectotype of Myriarthrus
cingulator and 12 syntypes, all Aachen, 2 specimens without label (coll. Foerster)
(ZSM). Italy: g', lectotype of Plectiscus flavopictus, Genoa (?), coll. Gravenhorst (ZI).
Netherlands: &, Hoge Veluwe, Kemperberg, 21.VII.1967 (coll. van Rossem); the
following 4 g' & 7 Q all from Ede, Sijsselt, trap, 3 & 15.IX.1970; 17, 22 & 30.V.1971,
9 & 21.V1.1972 (CJZ); a (genitalia, Pl. 1 Fig. 3), Ede, trap, 8—15.IX.1970 (coll.
van Rossem). No locality, 9, coll. Smits van Burgst (ELW).

Megastylus pumilio Foerster

& 2 Megastylus pumilio Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 105.

& Megastylus retroligatus Foerster, 1871, Verh. naturh. Rheinl. 28 : 105. New synonymy.

& Megastylus leptoderus Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 105. New synonymy.
& Megastylus pauxillus Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 105. New synonymy.

280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 7, 1974

Characteristics of the type specimens. — Megastylus pumilio. Male. Front wing 2.5
mm. Antenna 31 segments. Labels: a white label “& 31 gl. Aachen”. Lectotype label
added by me. There are two specimens on the pin; the left one is selected as the lecto-
type.

Megastylus retroligatus. Male. Front wing 2.7 mm. Labels: a white label “& + 25
gl. Altenberg 28.5". A blue rimmed box label “retroligatus Frst.”. Holotype labelled
by me: M. pumilio det. van R.

Megastylus leptoderus. Male. Front wing 2.4 mm. Labels: a white label “4.30 gl.
Boppard 24.4”. A blue rimmed box label “leptoderus Frst.’. Holotype labelled by me:
M. pumilio det. van R.

Megastylus pauxillus. Male. Front wing 2.4 mm. Labels: a white label “&.29 gl.
Aachen”. A blue rimmed box label “pauxillus Frst.”. Holotype labelled by me: M.
pumilio det. van R.

Comment. — A number of very small specimens (& 2.7—3.2 mm; 9 2.5—3.2 mm)
probably represent a third species, and one difficult to recognize. From the available
names for this taxon I have chosen M. pumilio as the most suitable (both sexes de-
scribed). I do not give full descriptions of both sexes but refer to the characters given
in the keys. Setting aside the characters of the body or wing length which are not very
reliable, the female is still rather clearly differentiated from the M. cruentator female
by the stout middle femur (index 3.6—3.8) (5.1—7.0 in M. cruentator). The male,
however, can only be separated from M. crzentator with unreliable characters, namely
its size and either entirely brown or black thorax.

Apart from M. pumilio, Foerster described another three species, viz. M. retroligatus,
M. leptoderus and M. pauxillus. Of each only one male is available. When studying
these three holotypes I had to conclude that the character given by Foerster for separating
these species, namely the position of the nervulus, is of no use. Besides, three different
females have yet to be found in the field.

Material examined. — Germany: 2 G', lectotype of M. pumilio and 1 syntype, Aachen
(coll. Foerster) (ZSM); 1 & & 2 9, Boppard, 24 & 26.IV (coll. Foerster) (ZSM); &',
holotype of M. retroligatus, Altenberg, 28.V (coll. Foerster) (ZSM); &, holotype of
M. leptoderus, Boppard, 24.IV (coll. Foerster) (ZSM); &, holotype of M. pauxillus,
Aachen (coll. Foerster) (ZSM); g, Aachen (coll. Foerster) (ZSM); &, Lousberg (bei
Aachen), 3.X (coll. Foerster) (ZSM); 2 g', Altenberg, 28.V (coll. Foerster) (ZSM).
Netherlands: 9, Arkel, 20.VIII.1967 (CJZ); &, Asperen, 1.VII.17967 (CJZ).

Megastylus rufipleuris (Foerster) new combination

9 Myriarthrus rufipleuris Foerster, 1871, Verh. naturh. Ver Rheinl. 28 : 103.

Characteristics of the holotype. — Myriarthrus rufipleuris. Female. Front wing 3.0
mm. Labels: a white label “2 39 gl Aachen”. A blue rimmed box label “rufipleuris
Erst”. Holotype labelled by me: M. rufipleuris det. van R.

Description of the female. — Front wing 3.0—4.0 mm. Head black. Labial palpi
short, maxillary palpi long; white to brownish. Clypeus strongly convex, yellow to
brown, with sparse, erect hairs. Face, yellow to brownish, polished, with suberect,
sparse hairs. Frons with weak antennal scrobes, polished. Inner orbits above antennal
sockets with broad white lining, which at most reaches the lower ocellus. Antennae with
about 39—40 short segments, strongly pilose.

G. VAN ROSSEM : The types of Megastylus 281

Thorax with rather characteristic colouring; pleuron 1, entire pronotum, episternum 2
(mesopleurae), including sternal part, all coxae and legs light yellowish to light brown.
Mesoscutum dark brown to black, except for streaks along prescutal sutures. Scutellum
light brown. Entire propodeum black. Thorax with sparse, suberect hairs. Wings hyaline.
Index of middle femur: 4.2—4.8 (4 specimens!), thus lying between M. cruentator and
M. pumilio.

Gaster with Ist and 2nd tergite relatively roughly alutaceous, following tergites more
polished. Apical segments in undamaged specimens with rather long widely spaced
hairs. Gaster fuscous with 3rd tergite lighter.

Male. — Unknown.

Comment. — The available material of this species is very scanty, not more than four
females. Nevertheless there is reason to believe that we are dealing with a separate taxon.
The females have the second tergite alutaceous and are thus differentiated from M.
flavopictus; they also have a conspicuous white lining on the upper inner orbit which
does not occur in any of pumilio or cruentator specimens I have so far seen. I have some
suspicion that M. rufipleuris females might be aberrant specimens of M. flavopictus,
with the second tergite alutaceous; the absence of males might support this view, but
especially in a group of species with so few true morphological characters. Males may be
hidden between those of other species (M. pumilio).

Material examined. — Germany: ®, holotype of Myriarthrus rufipleuris, Aachen
(coll. Foerster) (ZSM). Netherlands: 2 9, Ede, Sijsselt, 28.IX.1970 and 12.V.1971
(CJZ); 9, Giessenburg, 1.VIII.1968 (CJZ).

Subgenus Dicolus Foerster, 1868

Type species: D. insectator Foerster, 1871 = M. impressor Schigdte, 1838.
Subgenus Dicolus: Thompson, 1888, Opusc. Ent. 12 : 1311.

Megastylus (Dicolus) inpressor Schigdte

Q Megastylus impressor Schigdte, 1838, Rev. Zool. Cuv. 1 : 139.
Megastylus impressor: Schigdte, 1839, Mag. Zool. 2(1) : 5—6.

® Dicolus insectator Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 97. New synonymy.
Dicolus insectator: Townes, 1971, Mem. Amer. Ent. Inst. 17 : 205.

Characteristics of the type specimens. — M. impressor. Female. Front wing 5.0 mm.
Labels: a white, double rimmed label ‘‘9 2.5. 1834 Strandm. Drewsen”. Strandm. =
Strandmgllen, a locality near Skousborg (Sielland)*. The date in Schigdte’s paper (1839!)
is 21.V.1834*. There is no doubt about the originality of the specimen. A second printed
label: “Danmark ex coll. Schigdte”. Holotype labelled by the present author.

Dicolus insectator. Female. Front wing 4.5 mm. Labels: a white label “9. 33 gl.
Aachen’. A blue rimmed box label “insectator Frst.”. Holotype labelled by me.

Female. — Postanellus slender and long, index about 7.0—10.0. Hind tibia slightly
inflated in proximal part (Fig. 4). Hind femur somewhat inflated in proximal part.
Front tarsus of normal shape, basitarsus about 0.33—0.43 of front tibia (Fig. 6). In
fresh specimens head, except clypeus, black, polished. Pronotum with yellowish to

*) Information from Dr. Borge Petersen.

282 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 7, 1974

reddish parts, mesoscutum black, polished. Mesopleurae and scutellum reddish brown.
Propodeum black with conspicuous fossa in proximal part. Legs yellowish to reddish.
Gaster with proximal tergites alutaceous. First and second tergite and apex brown to
black, third and fourth (in part) tergite yellow to brown. Ovipositor, when not bent
downwards, concealed, thus causing confusion with the male.

Male. — Males are difficult to recognize notwithstanding some important characters:
the hind tibia is not inflated proximally and there is no constriction in the distal part as
in males of M. pectoralis (compare Figures 7 and 8). The hind femur is long and
slender, index 7.0—8.5 (some overlapping with M. cruentator). The postanellus is
long and slender, index 8.8—12.5. The propodeum shows the conspicuous transverse
impression in its 0.25—0.28 proximal part, typical for the subgenus Dicolus. The three
specimens studied show a vividly brown coloration, with clypeus, latero-ventral parts,
legs including coxae and spot on gaster, yellowish brown.

Material examined. — Denmark: © , holotype of M. impressor, Strandmellen,
2.V.1834, leg. Drewsen (coll. Schigdte) (MC); 9, Strandmollen, leg. Drewsen (coll.
Schigdte, not a type spec.) (MC); dg, Viborg, leg. Schigdte (coll. Schigdte) (MC).
Germany: 9, holotype of Dicolus insectator, Aachen (coll. Foerster) (ZSM); 9, Nd
Spessart, Lochmühle, 17—22.X.1971, trap (coll. van Rossem). Netherlands: &, Planken
Wambuis (near Ede), 5.IX.1964 (coll. van Rossem); 9, Ede, 9.VI.1971, leg. Zwak-
hals (CJZ). No locality: @, Pt Ballon, 18.VIII.1961, 1100 m, leg. den Hoed (KWR);
d', leg. Schmiedeknecht (coll. Smits van Burgst) (ELW).

Megastylus (Dicolus) pectoralis Foerster

4 Q Dicolus pectoralis Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 97.

Characteristics of the type specimens. — Dicolus pectoralis. Female. Front wing 4.6
mm. Labels: a white label “©. 33 gl. Lousb. (= Lousberg, the type locality) 25.10”;
a white label (not original) “pectoralis 7 9 *)” in pencil; a label “Dicolus pectoralis
Foerst. 9 det. J. F. Aubert”; lectotype labelled by the present author. A second speci-
men, a male, undoubtedly a syntype, was labelled paralectotype.

Female. — The two characters of the subgenus Dicolus are well developed in M.
pectoralis female: namely the slender postanellus, index about 8.0, and the deep trans-
verse impression of the propodeum. Confusion with the subgenus Megastylus is out of
the question. The conspicuously inflated hind tibia distinguishes this species from two
other Dicolus, viz. excubitor and impressor, and Foerster quite rightly used this character
(Fig. 3). An interesting feature which this female shares with D. excubitor and the
doubtful D. subtiliventris, is the long regular pilosity of the antenna, not found in the
other Megastylus species discussed in this paper.

Male. — The only two available males were in Foerster’s set. These two show,
somewhat weakened, the characteristic shape of the hind tibia of the female. (Compare
Fig. 7). I think these males are M. pectoralis. The two specimens both have the face
entirely ivory-yellow with the inner orbit above the antennal sockets marked yellow, as
in the female. The extremely slender front and middle femora, indices between 7.4—
10.0, are very conspicuous which makes differentiation from Megastylus s.str. males easy.
The deep transverse propodeal impression of Dicolus is also present.

*) Probably not all female specimens are syntypes.

G. VAN ROSSEM : The types of Megastylus 283

Biology. — Considering that nothing is known about the biology of this species it is
perhaps worth noting that C. J. Zwakhals caught 4 females in his Malaise - Townes
trap on 1 November 1970 on the northern slope of a diluvial hill covered with mixed
wood (Fagus, Quercus, Pinus and some other components), about 75—100 years old.
Microclimate of the site was cool and rather moist. Townes (1971) notes for the
Microleptinae “moist habitats, such as cloud forests”.

Material examined. — Germany: 9, lectotype of Dicolus pectoralis, Lousberg (bei
Aachen), 25.X. (coll. Foerster) (ZSM); &, paralectotype of Dicolus pectoralis, Lous-
berg (bei Aachen), 25.X (coll. Foerster) (ZSM); 5 9, Lousberg (Aachen), respect.
2 9, 16.VIL.; 3.X.; 15.X.; 25.X. (coll. Foerster) (ZSM); & ©, Aachen (coll. Foer-
ster) (ZSM). Netherlands: 4 9, Ede, Sijsselt, 1.XI.1970, trap, leg. Zwakhals (CJZ).

Megastylus (Dicolus) subtiliventrie Foerster
Q Dicolus subtiliventris Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 97.

Characteristics of the type specimen. — Dicolus subtiliventris. Female. Front wing 3.1
mm. Labels: a white label “9, 31 gl. Lousb. (Lousberg bei Aachen, the type locality)
25.10”. À blue rimmed box label “subtiliventris Frst.”. Holotype labelled by the present
author.

Comment. — The treatment of this species must be tentative because material available
is limited, namely the holotype and only one corresponding specimen. The holotype and
the second specimen do not differ essentially from M. pectoralis. There are only two
points of difference, namely the smaller size and the absence of a broad yellow spot on
the frontal orbit droso-lateral of the antennal sockets (present in M. pectoralis), though
these characters are unhelpful in distinguishing it. I have at present treated M. subtiliven-
tris as a separate taxon, since a number of specimens of M. pectoralis from two quite
different localities all have the yellow spot, while both specimens of M. subtiliventris
also from two separate localities, do not. Finally it should be noted that part of Foerster’s
material of M. pectoralis and the holotype of M. subtiliventris were collected in the
same place on the same date!

Male. — Unknown.

Material examined. — Germany: 9, holotype of Dicolus subtiliventris, Lousberg (bei
Aachen) 25.X. (coll. Foerster) (ZSM); 9, Nd Spessart, Lochmiihle, 17-22.X.1971,
trap (coll. van Rossem).

Megastylus (Dicolus) excubitor Foerster

2 Dicolus excubitor Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 97.

Characteristics of the type specimen. — Dicolus excubitor. Female. Front wing 4.8
mm. Labels: a white label “©, 33 gl. Aachen”. A blue rimmed label “excubitor Frst”.
Holotype labelled by the present author.

Comment. — Of this taxon only the holotype was available. This specimen shows
relationships towards M. pectoralis, having the long pilose antenna and the slender
front basitarsus (about half as long as front tibia*) (Fig. 5), but at the same time the

*) Basitarsus about 0.57—0.59 of tibia in subtiliventris, 0.57 in pectoralis lectotype, 0.33 in impres-
sor lectotype.

284 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 7, 1974

differential character of the latter species is lacking, namely the inflated hind tibia.
Though the shape of the hind tibia has much in common with that of M. pectoralis and
some vague inflation could be suspected, the difference does not quite justify a synonymy
of M. excubitor with M. pectoralis. The solution of this question has to wait for further
material and for the present we shall have to accept Foerster’s differential character in
the hind tibia.

Male. — Unknown.

Material examined. — Germany: 9, holotype of Dicolus excubitor, Aachen (coll.
Foerster) (ZSM).

TYPE NOT IDENTIFIED

& Megastylus facialis Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 105.

Characteristics of the type specimen. — M. facialis. Male. Front wing 2.9 mm. Face,
including clypeus and oral parts ivory. Thorax entirely brown, except for a narrow
ivory-yellow rim on anterior margin of pronotum. All coxae and further parts of legs
light yellowish to brown, except for fuscous distal part of hind tibia. Index of postanel-
lus, 6.0. Index of hind femur, 5.3. Labels: a white label “&, 23 gl. Boppard 24.4”; a
blue rimmed box label “facialis Frst.”. Holotype labelled by the present author.

I have not succeeded in recognizing this holotype at the species level; certainly there
seems no reason to regard it as a separate taxon. The specimen has much in common
with M. pumilio although the true M. pumilio has a brown face. Definite identification
has to wait for further information with regard to e.g. M. rufipleuris and M. pumilio.

SPECIES INQUIRENDAE

Q Megastylus orbitator Schiodte, 1838, Rev. Zool. Cuv. 1 : 139 (type lost).

dg Myriarthrus aemulus Foerster, 1871, Verh. naturh. Ver. Rheinl. 28 : 103. The
orbita frontalis has a white lining in the type specimen. Foerster wrote: “Orbita frontalis
nicht weiss.” The specimen belongs to Megastylus flavopictus Gravenhorst.

I have not seen the type of the following species:

Miomeroides transsylvanicus Kiss, 1924, Verh. Mitt. Siebenbürg. Ver. Naturw.,
Hermannstadt 72—74 : 113. According to Townes (1971), a species of Megastylus.

Megastylus (Dicolus) hirticornis Strobl, 1903, Mitt. Nat. Ver. Steierm. 40 : 141.
Strobl writes: “Steht etwa neben swbtiliventris Frst., Thms., ist aber fast dreimal grösser
und auch durch die Färbung ausgezeichnet”. In all probability this is a specimen of M.
pectoralis.

Megastylus pleuralis Thomson, 1888, Opusc. Ent. 12 : 1313. Probably a specimen of
M. cruentator. |

ACKNOWLEDGEMENTS

For the loan of type material and specimens I am grateful to: B. Petersen, Universitetets Zoolo-
giske Museum, Copenhagen (MC); W. J. Pulawski, Muzeum Zoologiczne, Wroclaw (ZI); E.
Diller, Zoologische Sammlung des Bayerischen Staates, Miinchen (ZSM); K. W. R. Zwart,
Laboratorium voor Entomologie, Wageningen (ELW); C. J. Zwakhals, Arkel (CJZ) and K. W. R.
Zwart (KWR) (the former G. J. den Hoed coll.). I am also indebted to Forschungsinstitut Sencken-
berg, Frankfurt a/M for kind hospitality in recent years and for giving me opportunity to do col-
lecting work in the northern Spessart at their Aussenstelle Lochmühle bei Bieber (Pl. 1 Fig. 1). It
was there that I collected some Microleptinae which arose my first interest in this subfamily.

G. VAN ROSSEM : The types of Megastylus 285

REFERENCES
Foerster, A., 1868. — Synopsis der Familien und Gattungen der Ichneumonen. — Verh. naturh.
Ver. preuss. Rheinl. 25: 135—221.
, 1871. — Uebersicht der Gattungen und Arten der Familie der Plectiscoiden. — Verh.

naturh. Ver. preuss. Rheinl. 28: 71—123.
Gravenhorst, J. L. C., 1829. — Ichneumonologia Europae 2 (2): 978—989 (Plectiscus). Vratisla-

viae.

Haliday, A. H., in Curtis, 1837. — A guide to an arrangement of British Insects 2 (2 ed.).

Kiss, von Zilah, A., 1922-1924. — Beiträge zur Kenntnis der ungarischen und siebenbürgischen
Ichneumoniden (Schlupfwespen) Fauna. — Verh. Mitt. Siebenbürg. Ver. Naturw. Her-

mannstadt 72—74: 32—146.

Perkins, J. F., 1962. — On the type species of Foerster’s genera (Hym. Ichneum). — Bull. Br.
Mus. nat. Hist., Entomology 2 (8): 383—483.

Schigdte, J. C., 1838. — Ichneumonidarum, ad faunam Daniae pertinentium, genera et species

novae. — Rev. Zool., Soc. Cuvierienne 1: 139.

, 1839. — Ichneumonidarum, ad Faunam Daniae pertinentium genera et species novae,

descripsit. — Mag. Zool. D'anat. Comp. et de Pal. II: 1—..., Pl. 6—10.

Schmiedeknecht, O., 1911. — Opuscula Ichneumonologica. Plectiscina. 28 & 29: 2161—2271.

Strobl, P. G., 1903. — Ichneumoniden Steiermarks. — Mitt. Nat. Ver. f. Steierm. 40: 43—160.

Thomson, C. G., 1888. — Försök till gruppering af sligtet Plectiscus (Grav.). — Opuscula ento-
mologica 12: 1266—1318.

Townes, H., 1971. — The genera of Ichneumonidae, part 4. — Mem. Amer. Ent. Inst. 17: 1—372,
PI. 1—217. Ann Arbor.

Viereck, H. L., 1914. — Type species of the genera of Ichneumon flies. — U.S. Nat. Mus. Bull

83: 1—186.

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Plate 1. Fig. 1. Aussenstelle Lochmühle bei Bieber (Northern Spessart). Phot. van Rossem. Fig. 2,

Megastylus cruentator, ovipositor of lectotype (MC). Fig. 3, Megastylus flavopictus, genitalia male,

Ede, trap, 8-15.IX.1970 (coll. van R). Fig. 4, Megastylus cruentator, genitalia male, Ede, Sijsselt,
7.1X.1963 (coll. van R)

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* Een sterretje duidt een naam aan, nieuw voor de wetenschap.
* An asterisk denotes a name new to science.

Uit dit register zijn weggelaten de namen voorkomende in artikelen, die reeds zijn voorzien van
eigen registers, nl. die van van Lith (p. 101) en de Jong (p. 271); dat van Lieftinck (p. 224) is
nog eens in dít register opgenomen.

DIPTERA

apicalis 24
Cnemopogon 24
flavitarsis 23
Haplegis 23
Lipara 2 et seq.
lucens 3 et seq.
pullitarsis 5 et seq.
rufitarsis 5 et seq.
similis 4 et seq.

HEMIPTERA HOMOPTERA

acaenae 135
Acyrthosiphon 104, 106
*agastachyos 108
Aphis 108 et seq.
argentinaeradicis 131
*astragalina 112
*atromaculata 114
*baccharicola 116
*californica 140, 141
corylina 124
cyperi 140, 142
daviaulti 139
Ericobium 124
*falklandicus 136
*hybrida 140, 143
inflata 132
*lagacei 129
*magna 126
Masonaphis 124, 126
*Megourina 128
*mimuli 119
*mulini 121
*mulinicola 123
nabali 132
Nearctaphis 131
Neonasanovia 132, 133, 134
nigra 133
*ssp. nodulosa (T. verrucosa)

*sijpkensi 153
*ssp. subverrucosa (T. verrucosa)
[151
Trichocallis 142 et seq.
Thripsaphis 137 et seq.
Uroleucon 153
utahensis 140, 149
*vandenboschi 106
verrucosa 141, 149
zirnitsi 134

HYMENOPTERA

aemulus 284

alashanica 179

*alveolatus 49, 68, 98

Amegilla 180, 222
Anthophora 183, 212, 222
*apatelia 171, 178, 215
*apostasia 168, 196

arundinator 21
*aurifasciatus 43, 65, 98
avernus 51, 77, 98

badia 223

ssp. basilewskyi (P. latiannula-

[tus) 48, 52, 96, 99

benoiti 44, 51, 73, 98
bidentatus 46, 50, 81, 98
bizonatus 183

Bombus 183

bucconis 167, 178, 201
capensis 48, 51, 92, 99
cingulator 273

congolus 41, 53, 98

cruentator 273, 275, 276 et seq.
ssp. defector (B. mendax) 183
deiopea 171, 178, 212
Deltoptila 223

detritus 21

Dicolus 273, 281 et seq.
dilectus 47, 90, 99

{150 *disconota 175, 185

ossiannilssoni 140, 145

ssp. pacifica (T. ossiannilssoni)

ealae 43, 48, 64, 99
Elaphropoda 186, 219, 222

[140, 145 *empeyi 41, 58, 98

Pentamyzus 135, 136
*pentatrichopus 104
producta 139, 146
*scabra 141, 147

Emphoropsis 159, 160, 180
erusus 42, 62, 99
erythrostoma 273
excellens 160

287

excubitor 275, 281 et seq.
facialis 284
flavopictus 273, 275, 278 et seq.
*freetownensis 43, 48, 67, 99
fulgidus 44, 49, 69, 99
fulvipes 186
garambae 43, 62, 99
ghesquieri 47, 87, 99
*Habrophorula 158, 217, 218
Habropoda 158 et seq.
habropodae 199
hirticornis 284
hookeri 177, 199
Ichneumon 273
“imitatrix 170, 178, 203
impressor 275, 281 et seq.
*jacoti 45, 50, 77, 99
kohli 45, 74, 99
krishna 216
laboriosa 160
ssp. laevior (Ps. capensis) 94,
[99
latiannulatus 48, 51, 94, 99
leclercgi 41, 53, 98
legionarius 23
*leoninus 45, 72, 99
liparae 23
luctuosus 45, 75, 99
lusingae 46, 76, 99
madecassus 41, 56, 98
magrettii 186
*ssp. maiella (H. tainanicola)
[193
mediator 273
Megastylus 273, 276 et seq.
melas 23
mendax 183
Mimesa 41, 58, 99
mimetica 164, 174, 183
Miomeroides 284
moelleri 186
montana 186, 205
montezumia 223
Myriarthridea 273
Myriarthrus 273 et seq.
nigeriae 46, 85, 99
nubilipennis 218
*orbifrons 173, 182
*oweni 47, 51, 89, 99

288

*ssp. pallidus (P. bidentatus)
[46, 50, 83, 98

orbitator 284

patei 43, 63, 99

patellatus 42, 58, 98

paulisae 42, 49, 61, 99

pauscillus 92

pectoralis 275, 282 et seq.

pekinensis 163, 172, 179

*pelmata 169, 210

percarinata 219

phragmitidis 21

*plantifera 170, 208

Plectiscus 273

pleuralis 284

Polemon 23

Protomelissa 188, 199

Psen 41 et seq.

Psenulus 97, 99

pseudobomboides 183

Pteromalus 23

pumilio 273, 275, 279 et seq.

radoszkowskii 169, 177, 205

reticulosus 44, 50, 70, 99

rowlandi 166, 175, 188

ssp. rubrocaudatus (P. bidenta-
[tus) 47, 84, 98

rufipleuris 275, 280 et seq.

*rugifrons 46, 80, 99

saltitans 43, 63, 99

*sapobaensis 44, 70, 99

Scambus 21

silvaticus 41, 52, 98

sinensis 164, 173, 180

Stenomalina 23

stevensoni 45, 50, 75, 99

stuckenbergi 92

subtiliventris 275, 283 et seq.

sutepensis 168, 175, 198

tainanicola 166, 176, 190

tarsata 186

Tetrastichus 23

transsylvanicus 284

trevirus 51, 88, 99

tricolor 188

*tumidifrons 173, 182

turneri 42, 49, 59, 99, 165,
[174, 185

*uelleburgi 47, 91, 99

zonatula 179

LEPIDOPTERA

adenensis 241

ssp. ali (sertorius) 247

f. alioides (sertorius) 251
ssp. amenophis (doris) 240
*brunneiflava 34

ssp. carnea (orbifer) 256
Clupeosoma 28

ssp. daphne (doris) 240
doris 239

ssp. evanida (doris) 241
flaviplagialis 29

f. gavarniensis (sertorius) 251
ssp. gecko (osthelderi) 237
geron 238

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 7, 1974

f. gracilis (sertorius) 250

f.guadarramensis (sertorius)
[251

ssp. hermona (phlomidis) 235

f. hibiscae (sertorius) 250

ssp. hilaris (orbifer) 254

hyalina 36

*Hyalinarcha 35

*iebelealis 31

ssp. kiki (phlomidis) 271

ssp. lugens (orbifer) 255

*minor 33

Neogenes 28

nigropunctalis 34

notodontalis 36

orbifer 252

osthelderi 235

phlomidis 233

f. posttesselloides (orbifer) 254

*Pseudonoorda 32

rufistriatum 28

sertorius 246

Spialia 228 et seq.

ssp. struvei (geron) 238

f. tesselloides (orbifer) 254

ssp. therapne (sertorius) 248

Thesaurica 36

*triangularis 30

*Trigonorda 29

AVES

Dryobatus major 24
Parus caeruleus 24

_No. 4. F. Willemse, 1968. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov
and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl.
D.Fl. 25 —.

No. 5. C. A. W. Jeekel, 1970. — Nomenclator generum et familiarum Diplopodorum, 412 pp.
D.Fl. 100.—. à ;

No. 6. J. G. Betrem, 1971. — The African Campsomerinae (Hymenoptera, Scoliidae), 326 pp.,
| 47 figs., 6 pls. D.Fl. 120—.

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